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OSMANIA UNIVERSITY LIBRARY 

Call No. Sl7.$/(Sf4f/ Accession No. 3 
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HANDBOOKS OF AMERICAN NATURAL HISTORY 
ALBERT HAZEN WRIGHT, ADVISORY EDITOR 



Handbook of Frogs and Toads 

ALBERT HAZEN WRIGHT AND ANNA ALLEN WRIGHT 



HANDBOOK OF 

Frogs and Toads 

OF THE UNITED STATES AND CANADA 

BY ALBERT HAZEN WRIGHT 

PROFESSOR OF ZOOLOGY, EMERITUS, CORNELL UNIVERSITY 

AND ANNA ALLEN WRIGHT 




THIRD EDITION 



Comstock Publishing Company, Inc. 

ITHACA, .NEW YORK, 1949 



COPYRIGHT 1933, 1942, 1949 BY 
COMSTOCK PUBLISHING COMPANY, INC. 



All rights reserved. This book, or parts thereof, must 
not be reproduced in any form without permission in 
writing from the publisher, except by a reviewer who 
wishes to quote brief passages in a review of the book. 



PRINTED IN THE UNITED STATES OF AMERICA BY THE 
VAIL-BALLOU PRESS, INC., BINGHAMTON, NEW YORK 



This volume, meant to serve the public, the scientist, 
and the group treated, is dedicated to the four Amer- 
ican women who, in addition to serving the public 
and science generously, have in the last half-century 
contributed most notably to the study of this group: 

Mary Hewes Hinckley, 1845-1944 
Mary Cynthia Dickerson, 1866-1923 
Helen Dean King, 1869- 
Helen Thompson Gaige, 1889- 



Preface 



NINETY YEARS ago John Le Conte in the prefatory paragraphs to 
his "Descriptive Catalogue of the Ranina of the United States" (Gen., 
1856, p. 423) wrote: 

Before I begin, it is necessary to observe that all the Ranina which I have ever 
seen have more or less the power of changing color at will. The character of 
color, therefore, of so much moment in the description of many other animals, 
is here of very little value; for none of the marks dependent on it are constant. In 
consequence, it requires numerous specimens, living subjects and long study to 
produce any description that approaches perfection. 

How far I have succeeded in my attempt remains to be seen. I have been long 
and sedulously engaged in my researches. Every description has been made from 
living specimens. 

In conclusion (p. 431) he observed: 

These are all the species of this family of reptiles which I have been able to 
see in a living state. I hold it to be impossible for any one to make a correct de- 
scription of an animal which has the power of changing its color at will, unless 
he has it alive; and this power they all possess in a greater or less degree; when 
preserved in alcohol they fade. The difficulty of procuring descriptions made on 
the spot where the objects are found, has rendered this part of natural history as 
confused and unsatisfactory as the researches of botanists who draw up their 
accounts of plants from dried specimens. 

In all our portrayals we have sought to emphasize the living animal by 
photographs from life, by color descriptions from life, by journal notes from 
the field, and particularly by excerpts from others' work pertaining to live 
frogs. In many regions we have been transitory visitors and the resident 
naturalists can tell a more complete story. Our notes and the notes of 
others, which in the two earlier editions were merged under the heading 
"Notes," are here expanded and designated "Journal notes" and "Authorities' 



corner." 



We have the warmest feelings for the countless friends and strangers who 
have helped us at different times and in different places. We have seen most of 
the preserved materials of the important collections and we thank their cus- 
todians. An especial effort has been made to secure live material and many 



viii PREFACE 

people have contributed. The first group constitutes individuals at most of the 
large zoos of the country. These people offered to loan or in some instances 
gave us much needed material. In a similar spirit specialists or workers one 
hundred or more in many places and institutions served us in countless 
fashions. The supply houses and biological bureaus were very eager and will- 
ing to help. The list qf old students who have mailed or shipped us live frogs 
would be hard to compile. Many have contributed to these almost weekly 
surprises through many years. The limits of this book forbid such a list. 
When we recall that, in California colleges alone, we have some twelve to 
fifteen teachers of biology, old students, we realize the size of our obliga- 
tions. Throughout this book will appear the collectors, helpers, comrades 
our creditors. 

For the typing of this work we extend our thanks to Mrs. Katharine Kapp, 
Dr. Ann L. Dunham, Mrs. Frankie Culpepper Goerges, and Mrs. R. F. Darsie. 
Two associates who have helped us particularly are Professors W. J. Hamilton, 
Jr., and Edward C. Raney. We are also indebted to Cornell University for the 
Faculty Research Grant to help in typing, purchase of specimens, visits to 
museums, and trips for specimens. 

ALBERT HAZEN WRIGHT 
ANNA ALLEN WRIGHT 
Ithaca, New 
December, 



Contents 



PREFACE . ... . vii 

GENERAL ACCOUNT . . . . . . i 

Common names . i 

Present classifications . .... . 2 

Scientific name . .2 

Specific evaluation of our North American forms . 4 

Range 5 

Habitat . .... .10 

Size . . . . ... . 10 

General appearance . . 12 

Color . . . ... .12 

Structure . . .... 12 

Voice 12 

Vocal sacs . . . ... 14 

Color of throat . . . . . 18 

Enlarged tympana (males) . . 20 

Breeding sizes . 20 

Summary of secondary sexual characters . . 24 

Ovulation . .26 

Egg-laying process . . 31 

Eggs . ... 31 

Synopsis of eggs of United States frogs . . 36 

Tadpoles . . .46 

Synopsis of United States tadpoles . . . . .* 54 

Development and transformation . . 75 

Journal notes . .... 79 

Authorities' corner . ... -79 

KEYS . . 81 

Families . . - - . . . . . 81 

Ascaphidae ... . . . . . 81 

Scaphiopodidae ... 82 

Bufonidae 83 

Hylidae . . ... 87 

Leptodactylidae . . ... . . 94 

Ranidae ... ... 96 

Brevicipitidae . . 103 



x CONTENTS 

Page Number of 

Account Plate Map 

ACCOUNTS OF SPECIES 104 

Ribbed toads, Ascaphidae 104 

American bell toad, Ascaphus truei 105 no 105 

Spadefoots, Scaphiopodidae in 

Couch's spadefoot, Scaphiopus couchii in no 112 

Hammond's spadefoot, Scaphiopus hammondn hammondii 113 117 114 

Central Plains spadefoot, Scaphiopus hammondii bombifrons 116 117 114 
Great Basin spadefoot toad, Scaphiopus hammondii inter- 

montanus 120 120 114 

Spadefoot, Scaphiopus holbroofyi holbroo1(ii 123 125 124 

Key West spadefoot, Scaphiopus holbroo\n albus 127 128 124 

Hurter's spadefoot, Scaphiopus holbrootyi hurterii 130 130 124 

Toads, Bufonidae 135 

Colorado River toad, Bufo aharius 135 139 136 

American toad, Bufo amcricanus americanus 140 139 144 

Hudson Bay toad, Bufo americanus copei 143 147 144 

Northwestern toad, Bufo boreas boreas 146 147 144 

California toad, Bufo boreas halophilus 150 153 151 

Amargosa toad, Bufo boreas nelsoni 154 153 151 

Southern California toad, Bufo californicus 156 161 136 

Yosemite toad, Bufo canorus 160 161 151 

Great Plains toad, Bufo cognatus . 164 168 144 

Spade-footed toad, Bufo compactihs 167 168 136 

Little green toad, Bufo debilis 172 175 172 

Black toad, Bufo exsul 176 175 151 

Canadian toad, Bufo hemiophrys 180 183 144 

Sonoran toad, Bufo insidior 184 183 172 

Marine toad, Bufo marinus 188 193 144 

Spotted toad, Bufo punctatus 192 193 195 

Oak toad, Bufo quercicus 197 200 207 

Southern toad, Bufo terrestris 199 200 140 

Nebulous toad, Bufo valliccps 203 205 195 

Rocky Mountain toad, Bufo woodhoush woodhousii 206 205 207 

Fowler's toad, Bufo woodhousii fowleri 210 215 207 

Tree frogs, Hylidae 217 

Cricket frogs, Acris 217 

Cricket frog, Acris gryllus gryllus 217 221 218 

Cricket frog, Acris gryllus crepitans 220 221 218 

Chorus frogs, Pseudacris 229 

Mountain chorus frog, Pseudacris brachyphona 229 233 230 

Brimley's chorus frog, Pseudacris brimleyi 234 233 230 



CONTENTS xi 

Account Plate Map 

Swamp cricket frog, Pseudacns nigrtta mgrtta 236 239 237 

Clarke's chorus frog, Pscudacns ntgnta clartyi 240 239 267 

Eastern chorus frog, Pseudacns mgnta jenarum 244 249 237 

Northern striped tree trog, Pseudacns mgrita septentrionahs 250 249 267 

Striped tree frog, Pseudacns mgrita tnsenata 253 259 267 

Florida chorus frog, Pseudacns mgnta verrucosa 258 259 237 

Western chorus frog, Pseudacns occtdentalts 262 265 

Little chorus frog, Pseudacns ocular js 264 265 267 

Ornate chorus frog, Pseudacns ornata 268 273 230 

Strecker's ornate chorus frog, Pseudacns strecf^en 272 273 230 

Tree frogs, Hyla 281 

Anderson tree frog, Hyla andersonn 281 284 282 

Canyon tree trog, Hyla aremcolor 287 289 293 

Viosca's tree frog, Hyla avivoca 294 297 295 

Mexican tree frog, Hyla baud inn 298 297 293 

Creen tree frog, Hyla cineiea cineiea 303 307 303 

Miller's tree irog, Hyla cinerea evittata 308 307 303 

Peeper, Hyla c rucijet crucifer 311 316 312 

Florida peeper, Hyla oucijer bartramiana 315 316 312 

Pine woods tree Irog, Hyla fetnoralis 319 325 295 

Barker, Hyla giattosa 324 325 282 

Pacific tree frog, Hyla regilla 329 337 267 

Giant tree frog, Hyla septentrionahs 338 337 282 

Squirrel tree frog, Hyla squirella 341 345 342 

Common tree toad, Hyla vcrsicolor versicolor 344 345 293 

Cope's tree frog, Hyla versicolor chrysoscelis 347 348 293 

Dusky tree toad, Hyla veisicolor phaeocrypta 353 354 

Sonora tree frog, Hyla wnghtorum 360 348 267 

Robber frogs, Leptodactyhdae 366 

Mexican cliff trog, Eleutherodactylus augusti 366 369 

Texan cliff frog, Eleuthei odactylus latrans 368 375 371 

Ricord's frog, Eleutherodactylus ncordn plamrostns 377 369 371 

White-lipped frog, Leptodactylus labtalis 383 385 371 

Camp's frog, Syrrhophus cam pi 389 391 371 

Gaige's frog, Syrrhophus gaigeae 394 371 

Marnock's frog, Syrrhophus marnocfyi 395 391 371 

Frogs, Ranidae . 399 

Texas gopher frog, Rana areolata areolata 399 403 404 

Northern gopher frog, Rana areolata circulosa 402 403 404 

Oregon red-legged frog, Rana aurora aurora 410 415 411 

California red-legged frog, Rana aurora drayton'n 416 415 411 

California yellow-legged frog, Rana boyht boylii 419 425 420 



xii CONTENTS 

Account Plate Map 

Sierra Madre yellow-legged frog, Rana boylii muscosa . 424 425 420 

Sierra Nevada yellow-legged frog, Rana boylii sierrae . 429 433 420 

Florida gopher frog, Rana capito . 434 433 404 

Slater's frog, Rana cascadae . . 439 443 411 

Bullfrog, Rana catesbeiana . . ... 444 443 449 

Green frog, Rana clamitans . 450 453 451 

Nevada frog, Rana fishcri . 454 507 499 

Southern bullfrog, Rana grylio . . . 459 466 460 

River-swamp frog, Rana hecl(schen . . . 465 466 467 

Utah frog, Rana onca . 473 477 499 

Pickerel frog, Runa palustris .... 476 477 467 

Meadow frog, Rana pipiens pipiens . . .481 484 498 

Plain meadow frog, I Mutant J Rana pipiens burnsi 483 484 499 

Mottled meadow frog, [Phase] Rana pipiens \andiyohi 488 493 499 

Southern meadow frog, Rana pipiens sphenocephala . . 492 493 499 

Western spotted frog, Rana pretiosa pretiosa 520 528 451 

Nevada spotted frog, Rana pretiosa luteiventns 527 528 521 

Mink frog, Rana septentrionalis . 533 534 451 

Dusky gopher frog, Rana sevosa . . . 537 404 

Wood frog, Rana sylvatica sylvatica 540 534 541 

Northern wood frog, Rana sylvatica cantabrigensis 544 545 541 

Mexican frog, Rana tarahumarae 553 557 521 

Sphagnum frog, Rana uirgatipes . 556 557 460 

Narrow-mouth toads, Brevicipitidae . . 561 

Taylor's toad, Hypopachus cuneus 561 563 562 

Mitchell's narrow-mouth toad, Microhyla areolata 568 562 

Narrow-mouth toad, Microhyla carolinensis 571 579 562 

Taylor's microhyla, Microhyla mazatlanensis . 574 562 

Texas narrow-mouth toad, Microhyla olivacca . 578 579 562 

BIBLIOGRAPHY 5 8 4 

INDEX . ... 623 



Handbook of Frogs and Toads 



General Account 



TN THE TREATMENT of each species, the topical outline is as follows: 
JL Common names, Scientific name, Range, Habitat, Size, General appear- 
ance, Color, Structure, Voice, Breeding, Journal notes, and Authorities' corner. 

Common names: We of the United States and Canada have concerned our- 
selves little with distinctive names for the amphibians. Most of the common 
names of the salamanders and frogs are collective. Just as all salamanders are 
generally called water dogs or lizards, so also the frogs are known by a few 
common names (toads, frogs, or tree frogs). Some people use the word 
"peeper" indiscriminately for several small frogs that call in the early spring. 
Others call peepers lizards. 

Normally we expect common names to come from the people at large, but 
with amphibians and reptiles most of the common names in literature are 
really bookish names. Many are translations of the scientific names. The names 
may come from widely different sources, of which the following arc a few: 

(1) The person after whom the species is named. Example: Couch's spade- 
foot. Scaphiopus cotichii Baird. 

(2) The person who named the species. Example: Viosca's tree frog, Hyla 
avivoca Viosca. 

(3) The person who first collected it. Example: Taylor's toad. Hypopachus 
cuneus Cope. 

(4) A country. Examples: Canadian toad, American bell toad, Mexican 
toad. 

(5) A state or province. Examples: Sonora hyla, Winnipeg toad, California 
red-legged frog. 

(6) Habitats. Examples: River-swamp frog, pond frog, house frog, salt- 
marsh frog, canyon tree toad, desert tree toad, crayfish frog, gopher frog, wood 
frog, savanna cricket, cliff frog. 

(7) Habits. Examples: Chameleon tree frog, solitary spadefoot, grasshop- 
per frog. 

(8) Structural characters. Examples: Ribbed toad, narrow-mouthed toad, 
toothless frog, femoral hyla, thick-skinned frog. 

(9) Voice. Examples: Bell frog, screaming frog, pig frog, rattler, chorus 
frog, cricket frog. 

(10) Color. Examples: Three-lined tree frog, striped tree frog, ornate tree 
frog, green toad, cinereous hyla. 

(n) Seasons. Examples: Spring peeper, shad frog. 



2 HANDBOOK OF FROGS AND TOADS 

(12) Miscellaneous sources of many kinds. Folklore: example charming 
toad (legend says this toad, Bufo terrestris, turns your eye green). Use (bait) : 
example pickerel frog. Weather signs: example rain frog. Odor: example- 
mink frog. 

Present classifications: This work has primarily the purpose of presenting 
the living animal. Through our forty years of study and teaching we have 
realized the inadequacy of vertebrae, skull, pectoral girdle, sacral diapophyses, 
teeth, disks, and other structural characters (largely osteological) in the classi- 
fication of a smooth (scaleless) group. We early sought to work out our North 
American life histories to supplement this lack but doubt if we will ever have 
knowledge enough to undertake the task or to clutter up scientific literature 
with preliminary suggestions. One of the serious attempts in this country has 
been Dr. CJ. K. Noble's doctoral dissertation, The Phytogeny of the Salientia, 
wherein he attempts to employ musculature as a supplement to osteology, as 
has been done for birds and mammals. And his study will surely have to be 
considered in the final analysis. But many forms need to be canvassed and 
many life studies undertaken before we rely too implicitly on any set of mor- 
phological characters. In the past many morphological osteological criteria 
have been overworked and overemphasized, and we are in a state of judicial 
suspense awaiting a nonhasty synthesis. Therefore we have temporized by 
employing Stejneger and Harbour's check list with no particular emphasis of 
our own devising. 

Scientific name: Any consideration of the scientific name that an animal 
bears implies an understanding of the scheme of classification. All living 
things fall into two groups or kingdoms. The plants are treated in the science 
of botany, the animals in zoology. The animal kingdom has several major sub- 
divisions or phyla, the last being the Vertebrata (vertebrates). In the vertebrate 
phylum, the various classes are known as fishes (Pisces), amphibians (Am- 
phibia), reptiles (Reptilia), birds (At/es)^ and mammals (Mammalia). We 
designate the study of fishes as ichthyology, that of birds, ornithology, that 
of mammals, mammalogy, but we group together amphibians and reptiles in 
the science of herpctology. This merging of the two groups is in a measure 
due to our inability to designate infallible characters of separation. A fish has 
fins, a bird, feathers, a mammal, hair, but reptiles and amphibians have no one 
positively distinctive character. 

The amphibians, to which the order of frogs (Salientia or Ecaudata) be- 
longs, have been variously defined. Some fifteen to twenty characters have 
been employed. Most living amphibians have naked skin and a larval aquatic 
stage. Normally, as tadpoles or larvae, they breathe with gills air dissolved in 
the water; and as adults they breathe with lungs. Two of the membranes about 
a developing mammal are absent in amphibians. There are three living orders: 

Apoda (caecilians) are limbless, blind, and wormlike. None occur in the 
United States or Canada. 



GENERAL ACCOUNT 3 

Caudata (salamanders) arc, as adults, tailed. 

Salientia (frogs) are, as adults, tailless. 

Seven families of Salientia or ecaudate amphibians are represented by 99 
species or subspecies in the United States and Canada. Family names in zool- 
ogy and botany end in idae. These seven families with the number of species 
and subspecies in the United States and Canada are: 
/) Bell toads, Ascaphidae i species 

2) Spadefoots, Scaphiopodidae 7 species and subspecies 

j) Toads, Bufonidae 21 species and subspecies 

4) Tree toads, Hylidae 30 species and subspecies 

5) Robber frogs, Leptodactylidae 7 species 

6) Frogs, Ranidae 28 species, subspecies and phases 

7) Narrow-mouthed toads, Brevici- 5 species 

pitidae 

Some of these families are divided into subfamilies. The ending for sub- 
family names is inae. Thus, the true frogs considered in this work belong to 
the family Ranidae and the subfamily Raninae. 

The family is divided into genera and the genera into species. 

Ordinarily a scientific name consists of three parts: the first name is the 
generic name, the second the specific name, and the third the author or de- 
scriber who first gave the name. The specific Latin name serves as an adjective 
agreeing in gender and number with the generic name, which is treated as a 
noun. This is the binomial system of nomenclature. The meadow frog might 
serve as an example. It is called Rana pipiens Schreber. The generic name is 
written with a capital and the specific name with a small letter. If the species 
be divided into one or more subspecies or races, the name may consist of four 
parts, namely, genus, species, subspecies, and authority. Such a name is an 
example of trinomial nomenclature. The swamp cricket frogs, Pseudacris 
nigrita (Le Conte), may be divided into several subspecies written thus, Pseu- 
dacris nigrita nigrita (Le Conte); P. n. septentrionalis (Boulenger), etc. 
Notice that abbreviations may be employed after Pseudacris and nigrita have 
been spelled out once. 

CATEGORIES 

Kingdom Animal Subfamily Raninae 

Phylum Vertebrata Genus Rana 

Class Amphibia Species boylii 

Order Salientia Subspecies sierrae 

Family Ranidae Authority Camp 

Name: Rana boylii sierrae Camp 

[Note: Observe that we use in this edition another example, Rana boylii 
sierrae Camp. In the 1942 edition, we used Rana pipiens burnsi (Weed) with 



4 HANDBOOK OF FROGS AND TOADS 

a purpose. Never have we been queried by youthful or elder scholars on any 
other issue as on this example. We employed it to stimulate work; unlike 
most of the critics, we saw these frogs in the field. Never did we seriously hold 
it to be a valid form, but in a book of this sort evidence ought to be presented 
with not too much youthful certainty or elderly obstinacy.] 

Specific evaluation of our North American forms. About 70 of our 99 spe- 
cies or subspecies are, in our opinion, established on firm grounds. If all the 
recently described forms were as certain as Ascaphus truei Stejneger, Rana vir- 
gatipes Cope, R. septentrionalis Baird, R. catesbeiana Shaw, and R. hec1(scheri 
Wright, no hesitation would be manifest. 

We have included accounts of all the forms of Stejneger and Harbour's 
check list. The last species, R. hecl(scheri, we never would have dared to de- 
scribe from preserved specimens alone had we not first seen its queer tadpoles 
and heard its peculiar voice. Nevertheless, we frankly put it in the list of forms 
for which more evidence is needed. In other words, we need field studies and 
much more ample material on at least 29 forms. They may all be tenable, 
but calm judgment dictates that material alive and preserved, life histories, 
and other evidence be at hand before they rank with established forms like 
A. truei, Scaphiopus couchii, Bufo debilis, Hyla gratiosa, and R. virgatipes. 
The 29 forms that need more attention are as follows: 

/) Scaphiopus hammondii bombtfrons 

2) Scaphiopus hammondii intermontanus 

j) Scaphiopus holbrool(ii albtts 

4) Bufo boreas nelsoni 

5) Bufo calif ornicus 

6) Bufoexsul 

7) Bufo compactihs (of Southwest) 

8) Bufo hemiophrys 

9) Bufoinsidior 
w) Acris crepitans 

//) Pseudacris nigrita (6 subspecies) 

12) Pseudacris occidentalis (probably nonexistent) 

/^) Hyla cinerea evittata 

14) Hyla crucifer bartramiana 

75) Hyla wrightorum 

16) Hyla versicolor chrysoscelis 

77) Hyla versicolor phaeocrypta (avivoca may be the same) 

/) Eleutherodactylus augusti 

79) Syrrhophus campi 

20) Syrrhophus gaigeae 

21) Rana areolata (subspecies) 

22) Rana boylii (subspecies) 
Ranacascadae 



GENERAL ACCOUNT 5 

24) Rana pipiens burnsi (color phase) 

25) Rana pipicns tyndiyohi (color phase) 

26) Rana pipiens (subspecies of the check list) 

27) Rana sylvatica latiremis 

28) Microhyla areolata 

29) Microhyla mazatlanensis 

The 99 species and subspecies have been established on different criteria 
and degrees of variations. It is the age-old problem. What is a species or sub- 
species? Are similarities in measurements indicative of relationship or of the 
same environment (e.g., short mouths and short legs in northern forms in 
contrast to longer mouths and hind legs in southern forms) ? Are rugosities 
(Pseudacris, Acris, Microhyla, Scdphiopus) sufficient for species or subspecific 
distinctions? Are they inherent or are they peculiar to certain southern or 
southwestern areas ? 

Is the absence of spots (R. p. burnsi) in a form derivative from a spotted 
form (jR. pipiens) sufficient for a species, or is it a mutant, a recessive, or a 
metachroistic phase ? We were of the group who had doubts regarding R. p. 
burnsi and R. p. \andiyohi, yet they were two types found among thousands 
of live frogs. Had they come as a few specimens (preserved material) in a 
collection from distant countries, we might with less knowledge have de- 
scribed them and accepted them readily. 

The 29 forms listed above are some of the most interesting and engaging 
ones for future study, and extended comments upon each appear under the 
species accounts. 

Range: Maps i to 3 give life zones of geographical distribution and rainfall. 
Maps 4 to 37 show species distribution. The core of this information will be 
found in A Chec\ IJst of North American Amphibians and Reptiles by Leon- 
hard Stejneger and Thomas Barbour (Cambridge: Harvard University Press, 
1943). We have added our own records through the years and such records as 
we have found in some of the smaller collections. 

The ranges were compiled from dot maps of all records for each form. They 
outline the outer bounds of each form but can in no way give all ecological 
evaluations each critic might wish. They are meant to give the layman a mac- 
roscopic view of the range of a species or subspecies without portraying indi- 
vidual dot records. Sometimes the map unavoidably includes a state in which 
there is no published or known record. Our ranges whenever they enter Mex- 
ico must be interpreted as approximate. 

Geologic speculations: Many features must be weighed in a study of distri- 
bution. Tolerance of drought and of high or low temperatures, as well as the 
type of food and shelter available, must be considered. The respective ranges 
of tolerance of concentrations of salt, sulphur, borax, potash, or gypsum must 
be noted. The susceptibility of either tadpoles or adults may limit the range. 
Frogs are probably very responsive to the rock structure of their abodes, their 




8 



i JS 



I 



GENERAL ACCOUNT 7 

breeding spots, at least, being dependent upon the surface water of the region. 

Do Pscudacris (cricket frogs) dislike granitic and other igneous situations? 
There is a strange lack of records of these small frogs in eastern Canada on the 
crystalline Laurentian Shield, although one form extends above the Arctic 
Circle west of Hudson Bay. The widespread Pseudacris n. triscriata (Map 18) 
stops short at the lava plains of the Snake River in Idaho but extends a knob 
of occurrence into the corner of that state near Montana where Carboniferous 
and Algonkian rocks occur and into northeastern Utah where there is lime- 
stone. Look at the range of P. brachyphona (Map 16). Those frogs seem to 
enjoy the carboniferous Pennsylvanian rocks of western Pennsylvania, West 
Virginia, corners of Ohio and Kentucky the Allegheny Plateau. P. n. fen- 
arum (Map 17) of Carlisle, Pa., occurs in the vicinity of Brunswick shale and 
other shales and limestones. Do marine limestone and marine shell marl help 
produce the rougher skin otP.n. verrucosa (Map 17) ? Do those same marine 
limestones of Key West influence the color of Scaphiopus h. albus (Map 7), 
vary the Microhyla species (Map 37) of the region, or affect the ventral color 
of Rana pipiens? 

Look for a moment at Rana p. prctiosa and 12. p. luteiventris (Map 35). The 
latter tolerates the tertiary volcanic rocks of the Columbia River Plateau and 
ranges through the lava-infested Snake River Valley and its tributaries into 
northern Nevada, while R. p. prctiosa prefers the continental deposits in and 
west of the Willamette Valley of Oregon, crossing Washington and coming 
down, behind the 12. p. luteiventris range, through the area of the Bitterroot 
Mountains and Grand Tetons to the Wasatch formations of northeastern 
Utah. 

Does Hyla c. evittata live at the fringes of the range of H. c. cinerea? 
Around Washington, D.C., the first seems to be crowded onto the Cretaceous 
rock formations, leaving H. c. cinerea to the east on the marl, alum bluff, and 
limestones of the Miocene and on the alluvial sands. 

We would all like to know more about finding the delicate little Syrrhophus 
marnoctyi, which chooses soft soil under stone or log shelter for habitat. 
Around San Marcos and Helotes, Tex., are Austin chalk, Eagle Ford shale, 
Edwards or Fredericksburg limestone, and the limestone to clay of the Wash- 
ita group in close association. In Tom Green County, where this frog has 
been found, occur similar structures with the possibility of San Angelo sand- 
stone and Blaine gypsum added. In Green Gulch, Chisos Mountains, we again 
find Eagle Ford shales, Fredericksburg limestone, Chisos sandstone tuff, 
touches of Washita group, and varying amounts of Austin chalk in close 
proximity. Does the different combination help vary the frogs 9 structure and 
pattern a bit, giving us S. gaigeae? 

What does the robber frog (Eleutherodactylus latrans) like? Its distribu- 
tion is surprising. For years it was known only from the region of San An* 
tonio, Tex., and from Madera Canyon in the Santa Rita Mountains in 



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jo HANDBOOK OF FROGS AND TOADS 

Arizona, a separation of almost 1000 miles. It has recently been found in New 
Mexico near Carlsbad, a limestone and gypsum country, and is reported in 
Arizona north of Roosevelt Lake, from Parker Canyon, where there is also 
limestone. There are unusually good combinations of rock conditions around 
Helotes, Hondo, and Uvalde, Tex., to make desirable homesteads for this no- 
tional form. The breaks of the Balcones Fault occur at the front of the lime- 
stones of the Edwards plateau allowing water to come to the surface, and there 
is plentiful underground water. The several kinds of limestone conserve the 
water, and gypsum content easily washes out, leaving caves, cavities, and worn 
rock ledges readily available for shelter. 

"Distribution of the Flora and Fauna of Louisiana in Relation to Its Geol- 
ogy and Physiography" (La. Acad. Set., 8, 11-73 [Dec., 1944]) shows well the 
interlocking of habitats, and Percy Viosca, Jr., explains (p. 55) how two forms, 
Acris g. gryllus and A. g. crepitans, may live in one pond separated by an in- 
visible barrier, the one at the inlet where acid water from pinelands is present, 
the other at the outlet where the alkaline influence of the tide is felt. 

Similar intimate studies of particular spots must be made before we can 
understand the apparent separation into broken spots and the apparent meet- 
ing of ranges of closely related forms. 

Habitat: This topic usually refers to nonbreeding habitats, but at times 
illusions are made to breeding localities as well. 

Size: The phrasing we have employed is: adults 3^-6% inches (males, 

$0-156 mm.; females, 87-165 mm.). These are the measurements of the large 

Colorado River toad, Bufo alvarius Girard. They mean that breeding adults 

* from 3% to 6% inches in length of body from tip to snout to rear end 

1 1 . fy back of tbr m*. "i ^ y/& inches or 80 mm. is the smallest size at 

Ji' ma A v5 rhature, and 6% inches or 165 mm. is the largest size of any 

.measured female. Almost invariably the lower measurement in inches will 

be that of a male, and the maximum adult measurement, the size of a female. 

Jordan and Evermann considered the two killifishes, Heterandria formosa 
and Lucania ommata, the smallest vertebrates of North America. We have 
collected many of these tiny fish, but confidently pronounce the little chorus 
frog, Pseudacris ocularis, much smaller. Cuba, we understand, has a still 
smaller frog. 

Most of the seven families have extremes in size. The tree frog adults vary 
from %o inch (11.5 mm.) in the little chorus frog (P. ocularis) to 5% inches 
(130 mm.) in the Key West tree frog (Hyla septentrionalis). The robber frog 
adults vary from % inch (15 mm.) in Camp's frog (Syrrhophus campi) to 3% 
inches (90 mm.) in the Texas cliff frog (Eleutherodactylus latrans). The adult 
toads vary from % inch (19 mm.) in the oak toad (Bufo qucrcicus) to 6% 
inches (165 mm.) in the Colorado River toad (B. alvarius). The adult spade- 
foots vary from i% inches (37.5 mm.) in Hammond's spadefoot (Scaphiopus 
A, hammondii) to 2% inches (72 mm.) in Holbrookes spadefoot (S. holbroo\ii 



M HANDBOOK OF PROGS AND TOADS 

holbroo\ii). The adult frogs vary from i% inches (36 mm.) in the northern 
wood frog (Rana s. cantabrigensis) to 8 inches (200 mm.) in the bullfrog 
(R. catesbeiana) . Two groups have little variation in their extremes. The 
narrow-mouthed toads vary in adults from % inch (20 mm.) in the Texas 
narrow-mouthed toad (Microhyla olivacea) to i% inches (41 mm.) in Taylor's 
toad (Hypopachus cuneus). The one species of ribbed toads (Ascaphus truei) 
varies from i% to 2 inches (28-51 mm.) in length. 

General appearance: In most cases these accounts were written with a live 
specimen or specimens in hand. Each gives the form of body (habitus) of the 
animal, the color of the animal, and some of the other outstanding characters. 
Often the animal is compared to a closely related species or to the common 
type of the group. 

Color: A consistent effort has been made to secure a color description from 
live specimens mainly in the field or occasionally in the laboratory. Each sex is 
described. It must be remembered each description is of a particular specimen. 
The description follows the Ridgway code (2d ed.), and his spelling. Any de- 
scription not following this code is marked "non-Ridgway." 

Structure: This section is meant to supplement the characters given under 
"General appearance" or to add to characters used in the keys. Whereas "Gen- 
eral appearance" is written from living animals, "Structure" is added from 
examinations of preserved specimens and from published descriptions. Unless 
otherwise stated, all measurements in descriptions or keys are relative to the 
body length (represented by L.). "Structure" is written in the abbreviated 
form preferred in scientific descriptions. Very frequently the original descrip- 
tion appears in this section. 

Voice: Early travelers often commented on the frog music of our country. 
Witness the following: 

"There be also store of frogs, which in the spring time will chirp, and whistle 
like birds; there be also toads, that will creep to the top of trees, and sit there croak- 
ing, to the wonderment of strangers!" 

"To the stranger walking for the first time in these woods during the summer, 
this appears the land of enchantment; he hears a thousand noises, without being 
able to discern from whence or from what animal they proceed, but which are, in 
fact, the discordant notes of five different species of frogs!" 

Previous to my coming to this country, I recollect reading the foregoing passages, 
the first in a history of New England, published in London, in the year 1671; and 
the other in a similar production of a later date. 

Prepared as I was to hear something extraordinary from these animals, I confess 
the first frog concert I heard in America was o much beyond anything I could con- 
ceive of the powers of these musicians, that I was truly astonished. This perform- 
ance was al fresco, and took place on the night of the i8th instant [April 18, 1794, 
Philadelphia], in a large swamp, where there were at least ten thousand perform- 
ers; and I really believe not two exactly in the same pitch, if the octave can possibly 
admit of so many divisions or shades of semitones. An Hibernian musician, who, like 



GENERAL ACCOUNT 13 

myself, was present for the first time at this concert of antimusic, exclaimed, "Be- 
gorrah, but they stop out of tune to a nicety" 

I have been since informed by an amateur, who resided many years in this coun- 
try, and made this species of music his peculiar study that on these occasions the 
treble is performed by the tree-frogs, the smallest and most beautiful species; . . . 
their note is not unlike the chirp of a cricket: the next in size are our counter tenors; 
they have a note resembling the setting of a saw. A still larger species sing tenor; 
and the under part is supported by the bull-frogs; which arc as large as a man's foot, 
and bellow out the bass in a tone as loud and sonorous as that of the animal from 
which they take their name. William Priest, Travels, London, 1802, pp. 48-50. 

Doubtless all males have voices, yet one has to be careful in such records. 
One of the phases of Pacific Coast frogs and toads about which all workers 
have the least evidence is the voice and vocal sacs; and it is a striking fact that 
Rana boylii, R. aurora, R. pretiosa, and Bufo boreas have no vocal sacs appar- 
ent in preserved specimens or particularly apparent in live individuals. Are 
they like some eastern forms, namely, R. clamitans, R. catesbeiana, and JR. syl- 
vatica? 

The impression that Boulenger gives and the layman has, namely, that fe- 
males are mute, is not exactly the situation. Let anyone pick up a female soli- 
tary spadefoot (Scaphiopus holbrootyi) and squeeze it, and he might think 
he had a male. Those who know the species would recognize the difference 
between the male and the female voice. Or lay this same female on her back 
and stroke her belly, and she will speak vigorously, possibly not so strenuously 
as a male but nevertheless the voice is there. All spadefoot females can and do 
croak some. In the same way females of spadefoots, frogs, and some other 
groups can open the mouth and give a cry or scream note just as young bull- 
frogs have alarm notes as they go skipping to cover. Any field naturalist, if he 
has heard the peculiar cry of frogs and toads when in the grip of a snake or 
turtle, very well knows females can make themselves heard. The mercy cry 
of a toad or frog (like jR. catesbeiana, R. pipiens, R. p. sphcnocephala, R. gry- 
lio, or some Hylas) can be and is as likely to be that of a female as a male. 
Miss Dickerson is quite right in saying that females are not necessarily voice- 
less. Doubtless in time we will find many species with more or less vocal 
females. 

Some of the adjectives employed in describing American frog notes are bub- 
bling, weird, plaintive, hoarse, woeful, mournful, complaining, nasal, inces- 
sant, musical, pleasant, whistling, prolonged, mellow, tremulous, squawking, 
shrill, deafening, ventriloquial, peeping, metallic, resonant, twittering, loud, 
guttural, snoring, snorting, gurgling, clacking, explosive, grating, and sweet. 

Man has attempted to characterize the voices of frogs since Aristophanes, 
but with varying success. Voice has been one of our most valuable clues on life 
histories, yet we must say it is hard for us to describe a call as others do. One's 
vehicle of description may be figures of speech, or it may be musical, phonetic, 



14 HANDBOOK OF FROGS AND TOADS 

mechanical, graphic, or biological. Frog voices can be portrayed by tonal 
graphs. They are also presented on phonograph records, such as "Voices ol 
the Night" (Comstock Publishing Company, Inc., Ithaca, New York). 

Each species usually has its distinctive type of breeding song, yet each species 
may have several variations. Hyla gratiosa may bark in the high- long-leaved 
pines, but "coatbet" in the breeding pools. A peeper on its way to the pond 
might possibly call differently from one with the chorus. One might call dif- 
ferently in cold or warm weather, dry or rainy spells. The breeding song is 
usually typical, but one needs to beware of believing it is the same at all times, 
The main thing is to learn the quality of the song in order to be ready for its 
variations. Frogs can be as individualistic in this regard as are other animals. 

It is a rather remarkable fact that one of the most frequent observations of 
earlier travelers (100 or 150 years ago) in this country was on the frog music 
of North America, but in the scientific names none suggest voice except in the 
early spring peepers (cricket frogs or savanna crickets, Acris gryllus, A. crepi- 
tans; chorus frogs or swamp cricket frogs, Pseudacris) and in three of the larg- 
est frogs, 12. clamitans, R. grylio, and R. pipiens. In the same way common 
names suggesting voice are not frequent. None of the vociferous spadefoots, 
toadsyand narrow-mouthed toads have common names suggesting their 
voices. 
^Several Hylas and others have come to be recognized as rain indicators. 

If one knows the frog notes, he can in one night do more work on frog dis- 
tribution than he might otherwise do in years. In one favorable night of auto 
riding from Lakehurst through southern New Jersey we located many colo- 
nies of JR. virgatipes which might have escaped us entirely under other cir- 
cumstances. The first record of a new form usually comes from a voice record. 
We never would have found our first Microhyla near Richmond, Va., or 
H. andersonii and JR. virgatipes on an overnight camp near Everett Pond, 
N.C., without first hearing them. This was our method near San Antonio in 
1925, namely, to go along the roads and listen until midnight or later and 
then pursue the species newly heard or most commonly calling that night. 

The choruses of the Southeast are immense. Sometimes as many as five to 
eight species may be breeding in one place. At a distance it may be a perfect 
din but to wade among the performers one encounters a deafening concourse 
scarcely imaginable. Often we would have to devise methods of culling out 
predominant voices to catch others desired. Sometimes we would vary the 
pressure of our own ear openings or cup our ears to sift one particular call out 
of the chorus, e.g., Hyla fcmordis would drown out Bufo quercicus. 

Vocal sacs (Plate I) : Doubtless all the males of our eastern species of frogs 
do croak and most of them have vocal sacs. Both males and females occasion- 
ally open their mouths to cry when in distress from teasing, alarm, injury, or 
capture, but all croaking males of the United States and Canada keep their 
mouths closed whether in air or under water. 




Plate L Vocal sacs. i. Southern toad, Bttfo terrestris. 2. Oak toad, Bufo 
quercicus. 3. Northern gopher frog, Rana areolata circulosa. 4. Narrow-mouth 
toad, Microhyla carolinensis. 5. Spadefoot toad, Bufo compactilis. 6. Spadefoot, 
Scaphiopus holbroofyi holbroofyi. 7. Cricket frog, Acris gryllus gryllus. 8. Little 
chorus frog, Pseudacris ocularis. 9. Pine woods tree frog, Hyla jemoralis. 
10. Barker, Hyla gratiosa. n. Gopher frog, Rana capita. 12. Southern bullfrog, 
Rana grylio. 13. Sphagnum frog, Rana virgatipes. 



16 HANDBOOK OF FROGS AND TOADS 

Two of our students have made a study of vocal sacs. In 1931 Miss Rachel E. 
Field made a detailed portrayal (unpublished MS) of the vocal sacs of our 
eastern forms, and in 1935 Dr. C. C. Liu examined our American forms and 
made comparisons with world forms. Dr. Lui found the following: l 

Vocal sacs and vocal sac openings absent (5) : 
Ascaphtfs truei Rana boylii muscosa 

Bufo halophilus Rana b. sierrae 

Rana aurora aurora 

Median subgular internal sac and slitlike openings (17) : 
Scaphiopus couchii Bufo hemiophrys 

Scaphiopus hammondii Bufo marinus 

Scaphiopus holbrookji Bufo punctatus 

Bufo alvarius Bufo terrestris 

Bufo americanus Bufo valliceps 

Bufo boreas Syrrhophus campi 

Bufo calif ornicus Syrrhophus marnoc1(ii 

Bufo canorus Hyla femoralis 

Bufo cognatus 

Median subgular external sac and slitlike sac openings (24) : 
Bufo compactilis Hyla gratiosa 

Bufo debilis Hyla regilla 

Bufo fotvleri Hyla squirclla 

Bufo quercicus Hyla versicolor versicolor 

Bufo woodhousii Hyla v. chrysoscelis 

Acris gryllus Pseudacris brachyphona 

Hyla andersoni Pseudacris nigrita 

Hyla arenicolor Pseudacris ocularis 

Hyla avivoca Pseudacris ornata 

Hyla baudinii Pseudacris triseriata 

flyla cinerea Microhyla carolinensis 

'Hyla crucifer Microhyla texensis 

Median subgular sac internal and round sac openings (i) : 

Hyla septentrionalis 

Median subgular sac external and round vocal openings (o). 
Paired subgular sacs internal and slitlike openings (o). 
Paired subgular sacs external and round openings (o). 
Paired subgular sacs internal and round openings (i) : 

Rana palmipes 

Paired lateral sacs internal and slitlike openings (2) : 
Rana sylvatica latircmis Rana septentrionalis 



1 The names are those used by Dr. Liu in his thesis. 




Plate II. Throat differences in male and female, i. Cricket frog, Acris 
gryllus crepitans, male. 2. Cricket frog, A. g. crepitans, female. 3. Narrow- 
mouth toad, Microhyla carolinensis t male. 4. Narrow-mouth toad, M. carolinen- 
sis, female. 5. Eastern chorus frog, Pseudacris nigrita feriarum, female, 6. 
Eastern chorus frog, P. n. jeriarum, male. 7. Spadefoot toad, Bufo compactilis, 
male (rear throat with pleats or lappet). 8. Southern toad, Bufo terrestris, 
female. 9. Southern toad, Z?. terrestris, male. 10. Great Plains toad, Bufo 
cognatus, male (rear throat dark), u. Barker, Hyla gratiosa, male. 12. Barker, 
H. gratiosa, female. 13. River-swamp frog, Rana hect(scheri t female. 14. River- 
swamp frog, R. hccl^cheri, male. 



i8 HANDBOOK OF FROGS AND TOADS 

Paired lateral sacs internal and round openings (9) : 
Rana boylii boylii Rana prctiosa 

Rana catesbeiana Rana septentrionalis 

Rana grylio Rana sphenocephala 

Rana palustris Rana sylvatica 

Rana pipiens 

Paired lateral sacs external and round openings (5) : 
Rana aesopus Rana draytonii 

Rana areolata Rana montezumae 

Rana clamitans 

These two were careful workers, yet we wish more information on live 
males to answer such inquiries as the following: 

1. Why should Bufo boreas and B. b. halophilus be in different categories? 
Or Rana aurora and R. a. draytonii? Or JR. boylii and R. b. sierrae, R. b. mus- 
cosa? 

2. Why is Bufo cognatus out of the B. compactilis, B. debilis, B. quercicus 
class ? 

3. Why is Hyla femoralis unlike our other Hylids (except H. septentri- 
onalis) P 

4. Why is R. sylvatica unlike R. s. cantabrigensis or R. s. latiremis? 

Color of throat (Plate II) : In Ascaphus truei, Syrrhophus campi, S. mar- 
nocf(ii, Eleutherodactylus ricordii, E. latrans, Bufo b. borcas, B. b. halophilus, 
R. s. cantabrigensis, R. aurora, R. boylii, R. pretiosa, R. sylvatica, the color of 
the throat of the male is little different from that of the female. These are 
forms with no prominent vocal sacs. In R. palustris, R. pipiens, R. p. spheno- 
cephala, R. capito, and jR. areolata, forms with lateral sacs, only slight differ- 
ences occur in the throat color. In the narrow-mouths each sex may have a 
dark throat. Usually the male's throat is darker, but it is often difficult to use 
this criterion. Sometimes in Scaphiopus holbroo1(ii and S. couchii the differ- 
ence is not readily recognizable. In S. hammondii enough of the dark slaty or 
bluish cast appears in the male's throat to distinguish it. 

The pronounced differences in throat coloration come in the Hylidae (ex- 
cept Pseudacris ocularis), Bufonidae, and Ranidae. Only seven of the twenty- 
one forms of Rana have this marked difference. These tend to have yellows 
(sulphur yellow, maize yellow, sulphine yellow, oil yellow, lemon yellow, 
olive ocher, aniline yellow, primuline yellow, or barium yellow) in the males, 
and in the females lighter washes of yellow or white; e.g., the green frog (R. 
clamitans) male has a barium yellow throat, the female a white throat. If each 
has the same yellow the male may have dark spots or some other equally strik- 
ing character. 

The whole throat of some species of toads will be discolored or darker than 
the rest of the venter, or only the lower part of the throat may be thus colored, 
as in the sausage group (Bufo cognatus, B. quercicus, B, compactilis). 




Plate 111. Enlarged tympana in male Ranas (seven species), i. Green frog, 
R. clamitans, male. 2. Green frog, 7?. clamitans, female. 3. River-swamp frog, 
R. hcc^scheri, male. 4. River-swamp frog, R. hcc\scheri, female. 5. Mink frog, 
R. septentrionalis, male. 6. Mink frog, R. septentrionalis, female. 7. Sphagnum 
frog, R. virgatipes, male. 8. Sphagnum frog, R. virgatipes, female. 9. Nevada 
frog. R. fisheri, male. 10. Nevada frog, R. fisheri, female, n. Southern bullfrog, 
R. grylto, male. 12. Southern bullfrog, R. grylio, female. 13. Bullfrog, R. catesbci- 
ana t male. 14. Bullfrog, R. catcsbeiana, female. 



20 HANDBOOK OF FROGS AND TOADS 

In the Hylidae, Pseudacris (except P. ocularis), Acris, and Hyla have the 
throat dark and the female has the throat white or like the rest of the venter. 
In the group to which Hyla cinerea cinerea, H. c. evittata, H. gratiosa, and 
H. andersonii belong, the males have the whole throat or most of the throat 
dark or green, but the females have green on either side of the throat below 
and ahead of the angle of the jaw. In the two subspecies of H. cinerea females 
the color varies in degree of extension of green on sides of throat. In H. gra- 
tiosa females the green extension is more or less demarcated on the throat side 
with a prominent white border, and in H. andersonii this white border be- 
comes clearly defined in females. 

In general, males, being more active, are inclined to darker colors on the 
dorsum than the females. In a mated pair this is often quite noticeable, but 
even in those not mated the contrast may be very pronounced. This discrep- 
ancy is marked in early spring breeders but also obtains for later breeder's. So 
also old males incline more to self-color (witness Ran a, etc.) than the females, 
which may be more spotted. Or if a pattern be common to each, it will be 
bright in the female (e.g., Scaphiopus holbrootyi females may have yel- 
lowish or buffy stripes or spots on back and the males may be almost uni- 
form). 

Enlarged tympana, males (Plate III) : The seven species of frogs (Rana) in 
which the throats of the males may be differently colored from those of the 
females have even more marked differences in the tympana. These seven spe- 
cies first breed at about 41 mm. for R. virgatipes, 44 mm. for R. fisheri, 48 mm. 
for jR. scptentrionalis, 52 mm. for R. clamitans, 82 mm. for R. hecl(scheri and 
R* grylio, and 85 mm. for R. catesbeiana. 



BREEDING SIZES 
Name Males Females 



mm. mm* 
Ascaphidae 

Ascaphus truei 29-43 28-51 

Scaphiopodidae 

Scaphiopus h. bombifrons 38-56 43~56 

Scaphiopus h. intermontanus 4~59 42-63 

Scaphiopus h. hammondii 42-52 44~5 2 -5 

Scaphiopus h. hurteri 43-73 44~8 2 

Scaphiopus couchii 48-70 50-80 

Scaphiopus holbroo1(ii 54*72 



GENERAL ACCOUNT 



ax 



Name 



Males 



Females 



Bufonidae 






Bufo quercicus 


19-30 


2O-32 


Bufo debilis 


26-45 


31-52 


Bufo insidior 








Bufo punctatus 


40-68 


42-74 


Bufo a. copei 


40 


75 


Bufo w. jowleri 


41-74-5 


56-82 


Bufo b. nelsoni 

- -1TT7** *"*" 


42-68 


46-89 


Bufo terrestris 


42-82 


44-88 


Bufo calif ornicus 


44-58 


44-4,8 


Bufo txsul 


44-59 


46-65 


Bufo canorus 


46-64 


45-74 


Bufo cognatus 


47-95 


60-99 


Bufo compactilis 


52-78 


54-91 


Bufo a. americanus 


54-85 


56-110 


Bufo vaUiceps 


53-98 


54-125 


Bufo hemiophrys 


56-68 


56-80 


Bufo w. woodhousit 


56-99 


58-118 


Bufo b. boreas 


56-108 


60-110 


Bufo b. halophilus 


62-101 


60-116 


Bufo alvarius 


80-156 


87-165 


Bufo marinus 




-220 


Hylidae 






Acris g. gryllus 


15-29 


l6 ~33 


Acris g. crepitans 


17-30 


20-35 


Pseudacris ocularis 


11.5-15.5 


12-17.5 


Pseudacris n. scptentrionalis 


1Q -32 


19-35 


Pseudacris n. clartyi 


20-29 


21-31 


Pseudacris n. nigrita 


21-28 


22-30 


Pseudacris n. feriarum 


21-30 


22-35 


Pseudacris . triseriata 


21-32 


20-37-5 


Pseudacris n. verrucosa 


22 


3 


Pseudacris brimleyi 


24-28 


27-32-5 


Pseudacris brachyphona 


24-32 


27-34 


Pseudacris ornata 


25-35 


32-36 


Pseudacris strecJ^eri 


25-41 


32-48 


JHyla crucifer 


18-29 


20-33 


Hyla squirella 


23-36 


23-37 



Name 


Males 


Females 




mm. 


mm. 


Hyla femoralis 


24-37 


23-40 


Hyla wrightorum 


24-38 


24-42 


Hyla regilla 


25-48 


25-47 


Hyla v. phaeocrypta 


28P-33 


3 2?- 3 6 


Hyla avivoca 


28-39 


3 2 -49 


Hyla arenicolor 


29-53 


29-53 


Hyla andersonii 


30-4 1 


3N7 


Hyla v. versicolor 


32-5 1 


33-6o 


Hyla v. chrysoscelis 


36-43 


35-49 


Hyla c. evittata 


36-47 


32-47 


Hyla c. cinerea 


37-59 


4 I ~ 6 3 


Hyla baudinii 


44-71 


44-89 


Hyla septentrionalis 


46-75 


5 2 -5-95 


Hyla gratiosa 


49-68 


50-68 


Leptodactylidae 






Eleuthcrodactylus r. planirostris 


15-25 


1 7~3 


Syrrhophus cam pi 


J 5-23 


J5-25-5 


Syrrhophus gaigeae 


21 


28 


Syrrhophus marnoclyi 


21-33 


22-39.5 


Leptodactyltis labialis 


35? 


-49 


Eleutherodactylits latrans 


48-74 


57-90 


Ranidae 






Rana s. cantabrigensis 


29-50 


34-56 


Rana sylvatica 


34-60 


34-68 or 82 


Rana b. boylii 


39-^7 


40-75 


Rana virgatipes 


41-63 


41-66 


Rana fisheri 


44-64 


46-74 


Rana b. sierrac 


44-72 


48-84 


Rana onca 


44-68 


51-84 


\Rana a. aurora 


44-63 


52-87 


Rana p. pretiosa 


44-75 


46-95 


Rana p. hiteiventris 








Rana b. muscosa 


45 -6 


47-81 


Rana palustris 


46-64 


49-79 


Rana septentrionalis 


46-71 


48-76 



$ 




Plate IV. Secondary sexual characters, i. Enlarged thumb, single pad, Rana pipiens 
pipiens, male. 2. Horny excrescences on thumb and first finger, Bujo amcricanus ameri- 
canus, male. 3. Enlarged thumb, double pad, Rana boylii muscosa, male. 4. Enlarged 
forearm, Bufo boreas borcas, male. 5. Horny excrescences on thumb and first finger, 
Scaphiopus holbroo\ii holbrootyi, male. 6. Prepollex, Hyla septentrionalis. 7. Plaited 
lateral vocal sac, Rana arcolata circulosa, male. 8. Dimorphism in mating pair, male 
smaller, darker, female larger, lighter, Hyla crttctjcr crucijcr. 9. Horny excrescences on 
arm and fingers, Bujo alvanus, male. 10. Longitudinal throat plaits, Pseudacris nigrita 
]eriarum, male. n. Double throat sac, Hyla baudinu, male. 12. Apron over concealed 
lower throat sac, Bufo cognatus, male. 13. Tail and thumb pad, Ascaphus truei, male. 14. 
Excrescences on thumb pad and forearm, A. truei, male. 15. Anal tube, A. truci, female. 



HANDBOOK OF FROGS AND TOADS 



Name 


Males 


Females 




mm. 


mm. 


Rana p. sphenocephala 


49-78 


53-82 


Rana cascadae 


50-58 


5 2 -74 


Rana p. pipiens 


52-80 


52-102 


Rana clamitans 


52-95 


58-100 


Rana tarahumarae 


58 


-"3 


Rana a draytonii 


63-100 


58-136 


Rana capita 


68-ioi 


77-108 


Rana a. areolata 








Rana sevosa 


-84 


-92.5 


Rana grylio 


82-152 


85-161 


Rana hecT(scheri 


82-131 


102-131 


Rana catesbeiana 


85-180 


89-200 



Microhyla olivacea 
Microhyla carolinensis 
Microhyla areolata 
Hypopachus cuneus 



Brevicipitidac 



20-28 
20-30 
24-28 
28-37-5 



19-29.5 
22-36 
23-30 
36- 



SUMMARY OF SECONDARY SEXUAL CHARACTERS 

(Plate IV) 

A. Thumb enlarged in male (Ranidae and H. septentrionalis) 
B. Enlarged tympana (Rana clamitans group) 

C. Males have external vocal sacs; males 41-71 mm., females 41-76 
mm. (Rana virgatipes group: R. fisheri, R. septentrionalis t R. 
virgatipes) 

CC. Males without external vocal sacs; males first breeding at 82-85 
mm., females 82-89 mm., except JR. clamitans males 52-95 mm., 
females 58-100 mm. (Rana catesbeiana group : R. catesbeiana, 
R. clamitans, R. grylio, R. hecJ(scheri) 
BB. No enlarged tympana 

C. Base of thumb swollen but no prethumb expansion (prepollex) 
D. No external vocal sacs 

E. Thumb pad in two parts with oblique groove (Rana 
boylii, 3 subsp., and R. aurora group somewhat, 3 
forms) 



GENERAL ACCOUNT 25 

EE. Thumb pad or enlargement not in two parts; web 
of hind foot of male more or less convex (Rana syl- 
vatica group: R. s. sylvatica, R. s. cantabrigensis, 
R. p. pretiosa) 
DD. External lateral vocal sacs; webbing margin of hind foot 

not modified in male 
CC. Base of thumb enlarged with a striking prethumb expansion 

or prepollex (Hyla septentrionalis) 
AA. Thumbs not enlarged nor ball-like nor padlike at base 

B. Black or dark excrescences on first, second, and third fingers 

C. "Tail" in male; excrescences on breast and inner edge of fore- 
arm; fore limb and hind limb enlarged, and hind foot also in 
male (Ascaphus truet) 
CC. No "tail" nor excrescences on breast or forearm 

D. No median vocal sac or darkened throat in males, with 
heavy forearms (Bufo boreas group; B. b. boreas, Z?. b. 
halophilus) 
DD. Median vocal sac present in male 

E. Vocal sac in lower throat (in alcohol, apron or bib- 
like), darkened in male (Bufo cognatus or sausage 
group : B. cognatus, B. compactilis, B. quercicus) 
EE. Vocal sac not principally from lower throat 

F. Throat slightly discolored and finger ex- 
crescences not very prominent (Bufo punc- 
tatus group: B. punctatus, B. debilis) 
FF. Excrescences normal and throat discolored 
(Bufo americanus group: B. a. americanus, 
B. w. fowleri, B. terrestris, B. w. woodhousii, 
B. hemiophrys, B. valliceps) 
FFF. Arm broader in male; excrescences cover 
whole of upper fingers and first finger ex- 
crescence extends onto wrist (Bufo alvarius) 
FFFF. Excrescences on top of fingers, not inner 
edge primarily; throats little discolored 
(Scaphiopus group) 
BB. No black or dark excrescences on fingers 

C. Throat of males with median sac revealed in alcohol by 
wrinkles, folds, plaits, or darker color (Hylidae, Leptodacty- 
lidae) 
D. Throat not discolored in male or particularly unlike that 

of female (Leptodactylidae, 5 species) 

DD. Throat discolored in male or particularly unlike that 
of female (Hylidae: Acris, Pseudacris, Hyla) 



26 HANDBOOK OF FROGS AND TOADS 

E. Distinctly discolored (Acris, Pseudacris, some 

Hylas) 
EE. Not so distinctly colored in male (Hylas) 

F. Females with green on throat (Hyla cinerea 
group) 
G. Green edged inside by white line (Hyla 

gratiosa, H. andersonif) 
GG. Green not so edged (Hyla cinerea cinerea, 

H. c. evittata) 

FF. Females with throat like rest of venter (Hyla 
avivoca, H. arenicolor, H. crucifer, H. femo- 
ralis, H. squirella, H. versicolor) 
DDD. Throat discolored in male and female, less so in latter 

(Microhyla) 
CC. Males with paired vocal sacs below angles of jaw (H. baudi- 

nif) 

Ovulation (Plates V-IX) : The vast bulk of frog species are in the South 
and Southwest, the warmer climes. In the North activity begins with the going 
out of ice or the first warning of spring and moisture. Most of our texts have 
been written for the northeastern states where few species occur, and so many 
writers have concluded that all frogs breed in early spring because Rana syl- 
vatica, R. pipiens, and Hyla crucifer do. In the same way, many thought all 
salamanders breed in early spring because Ambystoma maculatttm, A. jeffer- 
sonianum, and A. tigrinum do. 

It is hard to classify species on one set of observations in one locality or re- 
gion. A species may be a short, early breeder in the northeastern states, e.g., 
Rana pipiens, yet lay eggs in July in Devil's River, Tex. A bullfrog may lay 
normally in June or July but, under peculiar circumstances, lay in February 
in San Antonio, Tex. A R. sphenocephala might breed early in February or 
almost any month in the South, and we might think it would be restricted in 
the North; yet we find it breeding in Wabash Valley, 111., in September. Even 
in the North Bufo americanus, B. jowleri, Scaphiopits holbrootyi, and R. 
clamitans stretch the ovulation periods a condition more normal in the South 
and Southwest. 

The spadefoots, toads, small tree frogs such as Acris and Pseudacris, some 
larger tree frogs (Hyla squirella, H. jemoralis, H. crucifer, H. regilla), and a 
few Ranas spawn in transient pools, impermanent situations,' roadside ditches, 
and temporary floodlands, and often the loss must be immense. In the North 
Rana sylvatica, R. s. cantabrigensis, and R. pipiens masses may be caught by 
the surface ice of spring freezes. Next to the losses in toads and spadefoots, 
we believe the species with small surface films suffer the most from complete 
drying up of pools, thus being hung up by evaporation above a pond's 
level. 



GENERAL ACCOUNT 




Plate V. Eggs: spadefoots, narrow-mouth toads (Scaphiopus, Microhyla). 
1,8. Hammond's spadefoot, 5. hammondii hammondti. 2,4. Spadcfoot, S. 
holbroot(it holbrooktt. 3. Couch's spadefoot, S. couchii. 5. Texas narrow-mouth 
toad, M. olivacea. 6,7. Narrow-mouth toad, M. carolinen$i$. 



28 



HANDBOOK OF FROGS AND TOADS 




Plate VI. Eggs: toads (Bujo). 1,2,5. Oak toad, B. qncrcicus. 3,8. Nebulous 
toad, B. valliceps. 4,10. Spotted toad, B. punctatus. 6. Southern toad, B. ter- 
restris. 7. American toad, B, americanus americanus. 9, Spadefoot toad, B. 
compactilis. 



GENERAL ACCOUNT 




Plate VII. Eggs: cricket frog, chorus frog, tree toad (Acris, Pscudacris, Hyla). 
i. Little chorus frog, P. ocularis. 2,5. Eastern chorus frog, P. nigrita feriarum. 
3. Cricket frog, A. gryllus gtyllus. 4. Clarke's chorus frog, P. n. clarfyi. 6. Striped 
tree frog, P. n. t riser iata. 7. Peeper, H. crucijer cructfer. 8. Squirrel tree frog, 
H. squire/la. 9. Green tree frog, H. cincrca. 10,13. P^ nc woods tree frog, H. 
femoralis. ir. Common tree toad, f/. versicolor versicolor. 12,14. Barker, H. 
gratiosa. 



HANDBOOK OF FROGS AND TOADS 




Plate VIII. Eggs: frogs (Ratta). i. Sphagnum frog, R. virgatipcs. 2. Gopher 
frog, R. capita. 3. Southern bullfrog, R. grylio. 4. Pickerel frog, R. palustris. 5. 
California yellow-legged frog, R. boyln boyhi. 6. Nevada spotted frog, R. pretiosa 
luteiventns. 7. Bullfrog, R. catesbeiana. 8. Wood frog, R. sylvatica sylvatica. 9. 
Green frog, R. cl ami tans. 



GENERAL ACCOUNT 31 

Egg-laying process: The egg forms may be summarized as follows: 

(1) Single eggs each singly ovulated after movement of pair, e.g., Hyla 
crucifer. 

(2) Single eggs, several strewn at one time followed by movement, e.g., 
several Hylas. 

(3) Single eggs laid as in (2) or small egg packets at surface, then move- 
ment by pair, e.g., Hyla cmerea. 

(4) Small packets of eggs ovulated at one time at the surface, movement by 
pair, e.g., Hyla versicolor. 

(5) Small packets of eggs ovulated at one time, no movement of pair and 
a resultant large film at the surface, e.g., Rana catesbetana. 

(6) Eggs laid in two rosarylike strings, e.g., Ascaphus truei. 

(7) Files (three- or two- or one-rowed), movement from place to place by 
pair, e.g., Bufos. 

(8) Bands, later swollen to cylinders, regular or irregular, like two films 
coalesced or two oviducal emissions merged; movement from bottom or base 
of stem to tip, e.g., Scaphioptts. 

(9) Balls, lumps, spheres, or plinths, merged oviducal emissions; mainte- 
nance of one position by pair during entire process or for many emissions, e.g., 
meadow frogs, gopher frogs, wood frogs, Pacific Coast frogs; Rana virgattpes 
and R. septentrionahs. 

The series is not so simple as it is given above because of movement or non- 
movement, surface or submerged laying, size of frog, egg complement, and 
season. 

Eggs: The species of the United States and Canada whose eggs are not re- 
corded are Scaphtoptis holbrootyi albus, Bttfo caltformcus, B. exsul, B. hemi- 
ophrys, Hyla avivoca, H. bauchnn, Syrrhophus campi> S. gatgeae, S. rnarnocfyi, 
Rana boyltt mitscosa, R. hecfyscheri, R. fishert, R. onca, Mtcrohyla areolata, 
M. mazatlanensis 15 in all. Breeding notes have been made but egg descrip- 
tions are lacking for Bttfo boreas boreas, B. debilis, Pseudacris ornata, P. nignta 
nigrita, P. n. septentnonahs, Hyla ctnerea evittata, H. wrightorum, H. versi- 
color chrysoscelis, Eletitherodactylus latrans, Rana areolata areolata, R. syl- 
vatica cantabrigensis, R. cascadae 12 in all. Of our 99 or more forms we thus 
have 27 species whose eggs and ovulation habits need more attention. In this 
country we know the eggs of about 70 species minutely. Of these almost 50 
species have been seen in the field. 

In the synopsis of egg characters we have drawn in part on the work of Dr. 
Tracy I. Storer for Rana aurora draytontt, R. boyht boylti t Bufo boreas ha- 
lophilus, Hyla arenicolor t and H. regilla. We have supplemented our observa- 
tions on Scaphioptts couchii and S. hammondii with Dr. Ortenburger's 
records. For Ascaphus and to some extent Syrrhophus marnocl(ii notes, we 
have used Mrs. Helen Thompson Gaige's collections and observations. E. R. 
Dunn's and R. F. Deckert's studies on Eleutherodactylus ricordii and Noble 
and Noble's work on Hyla andersonii have been incorporated in the synopsis. 



32 HANDBOOK OF FROGS AND TOADS 

Relatively, the egg vitelli of the robber frogs (Syrrhophus campi and S. 
marnoctyi, 3-4 mm. in diameter for yolk), in which the whole development 
occurs in the eggs, are the largest, with Ascaphus next. Actually, the largest 
vitelli are those of the bell toad, Ascaphus truei, in which yolks of 5 mm. occur. 
The smallest are those of the tiny swamp cricket frog, Pseudacris ocularis, with 
vitelli 0.6-0.8 mm. in diameter. 

The size of the adult or parent does not determine the size of the egg or, 
more strictly, its vitellus. The bullfrog, the largest, has vitelli 1.2-1.7 mm -> little 
larger than those of the narrow-mouthed toad, Microhyla carolinensis (1.0-1.2 
mm.) or of the tree toad, Hyla arenicolor, and smaller than those of Ricord's 
frog (Eleutherodactyltts ricordii plantrostris, 2 mm.). The bullfrog eggs are 
only 0.6-0.9 mm - Digger than those of our smallest species, Pseudacris ocularis. 
Within one group wide variation takes place. Why has the canyon tree toad 
(Hyla arenicolor) vitelli 1.8-2.4 mm., almost twice those of its relative, H. fe- 
moralis, 0.8-1.2 mm.? Or why should one of the smaller frogs, the wood frog 
Rana sylvatica, have eggs 1% times larger than the largest, the bullfrog? In 
the Northeast we concluded (in 1914) that the bullfrog had the smallest 
vitellus in proportion to its adult size and that Pseudacris n. triseriatus and 
Rana sylvatica had relatively the largest vitelli. In the Okefinokee region, 
Geo. (in 1929), we concluded that Bttfo quercicus, Pseudacris ocularis, P. n. 
nigrita, Acris grylltts, and Microhyla carolinensis were largest in vitelli rela- 
tively, and Rana grylio and 1?. catesbeiana were the smallest. 

The season of breeding for species in the North has its beginning and end 
clearly imrked. Each species needs four or five weeks, except Btifo a. amcri- 
cantis and Rana clamitans. The exceptions may require two or three months 
for ovulation. In the southeastern states, when once a species has begun, its 
season of breeding may extend throughout the summer or even into the early 
fall, depending upon the high crests of precipitation. These species, although 
of a swampy region, wait for the rains, and in this reliance on precipitation 
they suggest our desert species of Texas and Arizona. Those that do not begin 
until June have at least eight to ten weeks of ovulation. This minimum period 
for a species of the South is the maximum period for a northern form. Species 
such as R. p. sphenocephala, B. terrestris, and Acris gryttus, which begin early 
in the season, breed during 25-30 weeks of the year, if not longer, or from Feb- 
ruary to October or November. 

When one comes into a town in the Big Bend country of Texas or in the 
desert region of Arizona during a rain, after a dry period of six, nine, fif- 
teen, or more months, one wonders how it is that spadefoots and toads are 
ready that instant for their congress of mating and ovulation. We can only 
explain it by supposing that the ovulation period of these species extends over 
the whole spring and summer and that all individuals are not ready at one 
particular moment, those appearing being only a small part of the population. 
Perhaps the frogs aestivate or hibernate with life processes slow, and the eggs 



GENERAL ACCOUNT 33 

in this condition may linger almost ready for ovulation for a long period. 

In the mountainous regions of the Far West the season of ovulation is dic- 
tated by the short term of four or more months. Forms like Rana b. sierrae and 
Bujo canorus have a very short season for ovulation and tadpole development. 
If B. canorus did not come from a group of short larval development small 
tadpoles we would not be surprised at its wintering over as larvae, as must 
some 1?. b. sierrae. In the same way one wonders if some larvae of R. s. canta- 
brigensis in the North may not be occasionally forced into this same condi- 
tion. In the North ice goes out so late that ovulation and development periods 
are considerably restricted. 

The number ot eggs in a complement may vary from 6 in Syrrhophus campi 
and S. marnocfyi, or 35-51 in the bell toad, or 100 in the smallest species, 
Pseudacrts ocularis, to 20,000 in Rana catesbciana, the largest form. The range 
in tree frogs (Hylulae) is from 100 (P. oculans) to 2084 (Hyla gratiosa) ; in 
the toads (Bufonidae) from 610 (Bufo quercictis) to 16,500 (B. b. halophilus) ; 
in the frogs (Ranidae) from 349 (R. virgatipes) to 20,000 (R. catesbeiand) . 
The complements of the narrow-mouthed frog (Microhyla carolinensis) and 
the spadefoot (Scaphioptts holbroofyii) are, respectively, 869 and 2332. 

The eggs of eight species, Hyla cinerea, H. femoralis, H. verstcolor, Rana 
catesbeiana, R. clamitans, R. grylio, Microhyla carolinensis, and M. olivacea, 
float on the surface of the water; the eggs of 30 or more species (Acris t Pseuda- 
cris f Ascaphus, Scaphiopus, Bttfo, 14 species of Rana) are submerged. In 
northern or southeastern states no form with buoyant eggs lays before May 10, 
yet in Texas Microhyla ohvacea may breed in March or Rana catesbeiana t 
rarely, in mid-February. The 19 or 20 normally early breeders have submerged 
eggs. These eggs usually have firm jelly envelopes, except those of Pseudacns 
and Scaphiopus , which have a consistency intermediate between the firm jel- 
lies of early breeders and the loose surface films of late breeders. One form, 
M. carolinensis, although it lays at the surface, has the most beautifully dis- 
tinct, firm eggs of all the species considered, yet its Texan relative, M. olivacea, 
has indistinct loose-jellied eggs. 

We had eggs laid in camp and in the laboratory by mated pairs caught in 
the field. Later the eggs in the field were determined by these original checks. 
Whenever possible the process of egg laying was observed in the field. Two 
species, one of the North and one of the South, were identified by the positive 
elimination of all the other resident forms. 

The measurements and color descriptions are based on the fresh eggs of 
average adults, not of very young females. Later these forms were checked 
with preserved material. For the eggs with loose outer envelope the outer mar- 
gin is indicated by dots (Plate IX). In one species the vitelline membrane is 
far separated from the vitellus, and the space is indicated by cross hatching. 
A summary of the egg characters of each species follows in the accompanying 
synopsis (pp. 36-46). 



21 



I6o 




22 

T- 

> 
23 



I 



/ 



I6b 



9o 



12 



V / 



7o 



24 



25 






I7b 



9b 



7o 





26 







27o 



27b 



7b 



10 



15 




28 



Plate IX. Diagrams of eggs ty^2V>). From R. L. Livezey and A. H. Wright, Amencan 
Midland Naturalist, 37, 213-216 (1947). i. Eleutherodactyhts ncordii plamrostrts. 2. 
Ascaphus truei. 3. Scaphiopus couchn. 4. Scaphiopus hatnmondn hammondn, the only 
species of Scaphiopus with two envelopes. 5. Scaphtopus holbroofyi holbroofyi. 6. Scaphi- 
opus holbtoo\n hutteni. 7a,b. B/o alvarms. Note odd wedge-shaped vitelh in b, a com- 
mon condition in this species. 8. Bufo amcncanus americanus. 9a,b. Bufo boreas halo- 
philus. 10. Bufo calijormcus. n. Bujo cognatus. 12. Bufo compactilis. 13. Bufo pu net at us. 
The only Bufo that deposits eggs singly. 14. Bufo quetctcus. 15. Bufo tenestrts. i6a,b. 
Bujo valhceps. I7i,b. Bujo woodhousn jowlert. 18. //y/ andersomt. 19. //y/ atemcolor. 
20. //y/tf cmerca cineiea. 21. f/y/^ cmcijcr. 22. //y/ jemorahs. 23. Hy/0 gratiosa. Vitelhne 
capsule far removed from vitellus.* 24. f/y/j regtlla. 25. //y/a squtrella. 26. Hyla versicolor 
versicolor. 27a,b. -4rrw gryllus. Rarely laid as in b. 28. Pseudacris brachyphona. 



29 



31 



32o 



\ 



32b 



ZZc 



33 34 



35 



36 







\ 




45 












57 58 



29. Pseudacns mgnta clartyi 30. Pseudacns n jenarum. 31. Pseudacns n. septcntnonalis. 
32a,b,c. Pseudacns n. tnsenata. Figure a is probably the true appearance of the egg; c is 
after Smith's (1934) description; b is a composite of a and c. 33. Pseudacns n. venucosa. 
34. Pseudacns oculans. 35. Pseudacns ornata. 36. Pseudacns strec\en. 37. Rana aurora 
aurora. 38. Rana areolata ctrculosa. 39. Rana aurora dray to nit. 40. Rana boyln boy In. 
41. Rana b. stctrae. 42. Rana cascadae. 43. Rana clamitans. 44. Rana capito. 45. Rana 
catesbetana. 46. Rana gryho. 47. Rana palustns. 48 Rana pipiens ptpiens 49. Rana p. 
sphenocephala. 50. Rana pretiosa pretiosa. 51. Rana p. luteiventns. 52. Rana septen- 
trionalts. 53. Rana sylvattca sylvattca. 54. Rana s. cantabngensts. 55. Rana vtrgattpes. 
56. Microhyla carolmensis carohnensts. 57. Mtcrohyla c. olwacca. 58. Hypopachus cuneus. 
* No vitelhne capsules are shown, except in this figure, Hyla grattosa. 



HANDBOOK OF FROGS AND TOADS 



SYNOPSIS OF EGGS OF UNITED STATES FROGS 2 

A. Eggs unpigmcnted. 

a. Laid in water Rosary like string in circular mass, yolk 5 mm. in 
diameter, capsule 5 8 mm A tadpole stage. Egg complement 35-49. 
Season May to Sept. 11. Ascaphus truti 

aa. Laid on land or in moist situations. (Whole development within 
egg, no larval stage ) (No data for Eleuthtrodactylus latrans.) 
b. Egg complement small, 6-10. 

c. Complement 6 or 7. Yolk 3-3.5 mm., av 3 o. Season April to 
May. Ovarian evidence. Syrrbopbus campi 

cc. Complement fr-io, yolk about 4 mm. Season April to June or 
July. Ovarian evidence. Syrrhophus marnockii 

bb. Egg complement large, 11-15. (R. F. Deckert and Dunn.) Yolk 
i mm. in diameter, outer envelope eventually 4 mm. Season 
April to August. Elcuthcrodactylus ricordii planmstris 

PlateLXXVIII-5,7 

A A. Eggs pigmented (upper dark -pole and lower light pole). Laid in water. 

B. Eggs deposited singly 
a. Envelopes two. 

b Outer envelope, diameter 1.4-1.0 mm., inner envelope, diameter 
1. 1-1.6 mm., vitellus, diameter o 8-1 o mm., eggs brown above 
and cream below. Egg complement 941. Season June 10 to Aug. n. 

Hyla squinlla 
Plates VII-8, IX-i 5 

bb. Outer envelope, diameter 3.5-4 o mm.; inner envelope, diameter 
i.gr i.o mm., vitellus, i 1-1.4, vitellus dark brown and creamy 
white, strewn among sphagnum. Egg complement 800-1000. 
Season May i to July 10. Hyla andersonii 

aa. Envelope single. 

b. Envelope 1.3 mm. or more. 

c. Envelope 6.7 mm. more or less, loose, sticky, eggs single, 
tandemhke, often Bufo fashion or lattice work or irregular. 
Vitelh 1.3 mm., black above and white or gray below. Season 
April to July. Scaphtopus b. burterit 



R L. Livczcy and A H Wright, "Synoptic Key," The American Midland 
Natural* \t> 37, 179-111 (1947). 



GENERAL ACCOUNT 37 

cc. Envelope 1.3-5.0 mm.; not Bufo fashion, frequently single and 
irregularly arranged, 
d. Vitellus 0.9-1 8 mm. 

e. Vitelline membrane far from vitellus, appearing as inner 
envelope i. 6-1.0 mm., outer envelope loose, glutinous, 
indefinite in outline, 13-5 mm., vitellus 1.0-1.8 mm. 
Egg complement 1084. Season March 3 to Aug. n. 

Hyla grattosa 
Plates VII-n,i4, IX-i 3 ; LXIX-6 

ee. No inner envelope or appearance of one, envelope firm, 
indefinite in outline, 1.3-3.6 mm. 

f. Vitellus 0.9-1 o mm., upper pole deep brown, buffy 
olive lower pole, rarely the appearance of an inner 
envelope. Egg complement 141. Season April 15 or 
earlier to Sept. i. Sometimes in masses. 

Acns . gryllus 

Plates VII -3, IX i 7 a,b. 

Acns &. crtpitam 

ff. Vitellus i o 1.3 mm., upper pole black, lower pole 
white, envelope 3 i 3.6 mm. Egg complement 1000 
to 3000. Season April to September. Bufo punctatus 

Plate VI-4, 10 

dd. Vitellus 1.8-1.4 mm., jelly envelope 3.8 5.0 mm. Season 
March i or earlier to July i or fall. Hyla arenicolor 

bb. Envelope 1.1-1.0 mm. 

c. Vitellus 0.6-0.8 mm. Egg complement 100. Season January to 
September. Pseudacns ocularts 

Plates VII i ; IX 34 

cc. Vitellus 091.1 mm. Egg complement 809 1000. Season March 
30 to May 15. Hyla c. cructfcr 

Plates VII-7, IX ii,LXVI 4 

BB. Eggs deposited in a mass, 
a. Egg mass, a surface film. 

b. Single egg envelope a truncated sphere, the flat surface above; 

egg vitellus black above and white below. 

c. Envelope 1.8-4.0 mm., outline always distinct, never lost in 

the mass, eggs firm and distinct like glass marbles, making a 

fine mosaic; vitellus i .0-1.2. mm. Egg complement 869. Season 

March to Sept. 3. Mtcrohyla carohnensis 

Plates V-6,7; IX- 5 6; CXXV- 5 



38 HANDBOOK OF FROGS AND TOADS 

cc. Single envelope 1.5-1.0 mm., vitellus i.o mm. Egg mass a loose raft. 
Egg complement 700. Season April to October. Hyfofacbus cuncus 

bb. Egg envelope outline indistinct, more or less merged in the jelly mass; 
jelly glutinous, envelope not a truncated sphere, 
c. Egg brown above, cream or yellow below. 

d. Egg packets small, masses seldom if ever over 2.0 sq. in. (115 sq. 

cm.), or 4 X 5 in. in diameter (10 X 1^.5 cm.). 

e. Inner envelope large, 1.1-3 .4 mm. ; outer envelope 3 .1-5 .o mm. ; 

vitellus 0.8-1.6 mm. Egg complement 343-500. Season May 19 

to Aug. n. Hyla c. cincrca 

Plates VII -9, IX-io 

ee. Inner envelope small, 1.4-1 o mm., outer envelope 4-8 mm. 
f. Packets small, seldom over 30-40 eggs; vitellus 1.1-1.1 mm. 
Egg complement 1803. Season May 10 to Aug. 13. 

Hyla v, wrsicolor 
Plates VII-n; IX-i6; LXXIII- 5 

ff. Packets large, sometimes 100-115 eggs, vitellus 0.8-1.1 mm., 
av. 0.95 mm. Egg complement 708. Season April i to Aug. n. 

Hyla femoralis 
Plates VII -10,13; IX-ii; LXVIII-4 

cc. Eggs black above and white below. 

d. Egg mass small; outer envelope loose, irregular, does not look 
like a mosaic; somewhat merged, inner envelope possibly present 
close to vitellus, vitellus 0.8-0.9 mm - Outer envelope 1.8-3.0 
mm., not truncate. Egg complement 645. Season March 15 to 
September. Mtcrohyla oltvacca 

Plates V- 5 ;CXXVI- 3 

dd. Egg packets large, loose, glutinous films, 35-675 sq. in. (118-3711 
sq. cm.). (Suspect R. heckschen is in this class.) 
e. Inner envelope absent, vitellus 1.1-1.7 mm., egg mass 144-675 
sq in (900-3711 sq. cm.) in area, or n X M in. (30 X 61 cm.) 
in diameter, egg masses among brush around the edge of ponds 
encircling Pontca'cna-likc vegetation in mid-pond. Egg comple- 
ment 10,000-10,000. Season March 15 to Nov. 8. 

Rana cattsbtiana 
Plates VIII-7, IX-4 5 ; XCIV-6 

ec. Inner envelope present, 1.8-4.0 mm., vitellus 1.4-1.0 mm. 

f. Egg mass seldom i sq. ft. (35-144 sq. in. or 118-900 cm.) 
in area, or 5 X 7-1*. in. in diameter; usually around edge of 
ponds, inner envelope elliptic, pynform, or circular, av. 



GENERAL ACCOUNT 39 

3.05 mm.; vitellus 1.4-1.8 mm., mode 1.4 mm., av. 1.5 
mm. Egg complement 1451-4000. Season May 13 to 
Aug. n. Rana clamttans 

Plates VIII- 9 ; IX- 4 j; XCV~4 

ff. Egg mass over i sq. ft. in area (144-188 sq. in. or 900- 
1800 sq. cm.), or n X n in. to n X 15 in. in diameter; 
usually in mid-pond, inner envelope av. 3.45 mm.; vitel- 
lus 1.4-1 o mm., mode 1.8 mm., av. 17 mm. Egg comple- 
ment 8000-15,000. Season March 4 to Sept. 15. 

Rana ffylio 
Plates VIIL 3, IX-46 
aa. Egg mass submerged, 
b. Eggs in files or bands. 

c. Eggs laid in bands which soon become loose cylinders extending 
along plant stems or grass blades. 

d. Envelope 1.5-5.6 mm., vitellus 1.4-1.0 mm., vitelh throughout 
jelly cylinder, if stalkhke on almost imperceptible short stalks. 
Egg complements 1000-3000. 

e. Envelope 3 8-5.6 mm., vitellus 1.4-10 Egg complement 
1331. Season March or earlier to October. 

Scaphtopus b. bolbrookii 
Plates V-i, 4 ,IX-5, XXI 5 

ee. Envelope 1.5-3.5 mm. Vitellus 1.4 1.6 mm. Egg comple- 
ment 1000-3000. Season April to August. Scaphtopus couchtt 

Plate XVII 5 

dd. Envelope 1.5 i.o mm., vitellus i 0-1.6, eggs in bands or cylin- 
ders on ends of grass stems, along vegetation stems, etc. Eggs 
near periphery of cylinder more or less on stalks 5 10 mm. long. 
Egg complement 1000-1000. Season mid-February to August. 

Scaphtopus b. bammondtt 
PlatcV-i,8, XVIII 7 

cc. Eggs in files. 

d. Files short (4-10 mm. in length), 4-8 eggs in short bcadhkc 
chain or bar, or many such files radiating from one focus, vi- 
tellus 0.8-1.0 mm., tube diameter 1.1 1.4 mm. Egg complement 
610, 766. Season April i to Sept. 5. Bufo qucrctcus 

Plates VI i, 1,5, IX- 14, XL -5 

dd. Files long (several feet in length or often a meter or more long), 
c. Vitellus i. 0-1.6 mm., tube diameter 1.8-4.6 mm. 
f. Inner tube absent. 



4 o HANDBOOK OF FROGS AND TOADS 

g. Envelope more than 5 mm. in diameter (5-6 mm.), firm, distinct; vi- 
tellus 1. 1-1.6 mm., black above, white or gray below; 35 eggs in 30 
mm. (i 3/16 in.). Season May and June. Bufo cahfornicus 

gg. Envelope less than 5.0 mm. in diameter (1.8-4.6 mm.). 

h. Envelope i 6-4.6 mm., distinct and firm; vitelli crowded in the 
files at first in double row, later more spread out but still crowded 
although at times in single row, 11 15 eggs in 30 mm. (i 3/16 
in.). Season March to May. B. woodhousii 

i. Egg complement smaller, 5000-10,000; egg envelopes possibly 
smaller. Eastern Missouri eastward. Season April to mid- August. 

B, w. fowleri 
Plate IX-i 7 a,b 

11. Egg complement larger to 15,000, western Missouri westward. 
Season March to July. B w. woodhousii 

hh Envelopes i 4 or less (i 8-1.4 mm ) 

i. Envelope slightly scalloped in appearance; tube tightly coiled; 
1.8 1.4 mm., narrow, vitelli crowded, vitelh grayish or greenish 
brown above, cartridge buff or sulphur yellow below, 14-30 
eggs in 30 mm., vitcllus i 1-1.6 mm. Season May i to July 10. 

Bufo compacttlts 
Plates VI- 9 , XXXIII-4 

li. Envelope not scalloped, rather loose but distinct in outline, 
vitelli black or deep brown above, tan or white below, i.r- 
1.7 mm in diameter Season March to Aug 15. Egg comple- 
ment 7500-8000. Bufo alvanus 

ff . Inner and outer tubes present (i in number), 
g. Partitions apparent between eggs. 

h. Outer envelope distinctly scalloped; envelopes laminated, outer en- 
velope 1.7 mm. at emargmation, i.o mm. at widest diameter if in 
single row, or i.o mm. and 1.6 mm. if eggs in double row; inner 
envelope 1.6 mm. Vitellus 1.1 mm , black above, white below, 
complement to 10,000. Season April to September. B. cognatus 

hh. Outer envelope not scalloped, 3 4-4.0 mm. in diameter, usually not 
two rows, inner tube distinct 1.6-1.1 mm.; vitellus 1.0-1.4 mm. 
black or dark brown above, white below, 18-10 eggs in 30 mm. 
(i 3/16 in.). Egg complement 4000-10,000. Season March to late 
July. Bufo a. amencanus 

Plates VI-7, IX-8; XXV-6 



GENERAL ACCOUNT 41 

gg. No partitions between each individual egg. 

h. Eggs in a single row within the jelly tube. Outer en- 
velope 1.6-4.6 mm.; inner tube close to the outer 
tube, inner tube 1.1-3.4 mm.; outer tube inclined to 
be slightly scalloped, distinct space between eggs; 
vitelli i. 3-1. 4 mm.; 7-8 eggs in 30 mm. (i 3/16 in.). 
Season last of February to September. Bufo temstrts 

Plates VI-6;IX-i 5 

hh. Eggs usually in a double row within jelly tube, 
more crowded. 

i. Vitelli small, 1.1 mm., outer envelope small, 1.8 - 
3.1 mm., purplish black and white, inner envelope 
1.6 mm., close to outer tube, 10 eggs in 30 mm. 
(single row), or 15-17 eggs in 30 mm. (double 
row), rarely single eggs. Season March to Aug. 15. 

Bufo valltceps 
Plate VI 3,8 

ii. Vitellus larger, 1.50-1 7 mm , black and white or 
cream, outer envelope loose but distinct, large, 
4 9-5.3 mm. in diameter, vitelli double- or triple- 
rowed, inner tube 3.5-3.8 mm , not very close to 
outer tube (boreas group). 

j. Vitelli larger, 1.65-1.75 mm Season January to 
July. Bufo boreas halophtlus 

)j. Same, except vitelli average smaller though 
their range is 1.5-1.75 mm. Season March to 
September. Bufo b. boreas 

bb. Eggs an irregular mas>s, jelly loose, envelope 3.1 6.7 mm., vitellus 
black above and white below. 

c. Single envelope 3.1-3.6 mm., vitellus j a 1.3 mm , egg mass sub- 
merged film on bottom or loosely strewn on bottom. Season April 
to September. Bufo punctatus 

Plate VI- 4,10 

cc. Single envelope 6.3 mm., vitellus 1.3 mm., egg mass various pat- 
terns, single, tandemlike, Bufo-hkc, lattice work, etc., at or near 
surface attached to vegetation. Season April to June. 

Scaphiopus h. hurt mi 



42 HANDBOOK OF FROGS AND TOADS 

bbb. Eggs in lumps. 

c. Egg mass a firm regular cluster. 

d. One jelly envelope. (At times R. syhatica, R. s.cantabngtnsis,nn& 
R. p. pntiosa appear in this category.) 

e. Egg mass a plinth or sometimes globular, firm. No indistinct 
inner envelope. 

f. Vitellus 1.4-1.8 mm., black above, sulphur or primrose 
yellow below, envelope 4.9-6.9 mm., av. 5.4 mm., eggs 
farther apart than in R. ptpicns or R. p. sphcnocephala; egg 
complement 350-500 eggs, egg mass 1.5 X i to 3 X 4 in. 
New Jersey, southward. Season June 11 to Aug. n. 

Rana virgatipes 
Plates VHI-i,IX-55 

ff. Vitellus 1.8 11 mm., black above, light tan below; en- 
velope 5-71 mm., av. 63, egg complement 1000-3000 
eggs Egg mass 3 x 3 to 8 x 6 in. Northern Nevada north- 
ward and westward. Season March to May. 

R. prctiosa lutcivcntns 
Plate VIII -6 

ec. Egg mass a sphere, inner envelope indistinct. (See dd. Two 
or three jelly envelopes R s syhattca, R. s. cantabrtgensts, R. 
p. prettosa.} 

dd. More than one jelly envelope, 
c. Two envelopes. 

f. Egg mass a sphere 1^-4 in. (6 35-10 cm.) in diameter, con- 
taming 1000-3000 eggs, outer envelope distinct, 
g Eggs black above and white below, inner envelope ap- 
parently absent, slightly evident under lens, 3.6-5.8 
mm., av. 3.8, outer envelope 5.1-9.4 mm., av. 6.4, vi- 
tellus 1.8-1 4 mm., av. 1.9. Egg complement 1000-3000. 
Season March 19 to May i. Rana s. sylvattca 

Plates VIII-8, IX-53, CXIX~5 

[h. Is R. s cantabngcnsis thus: vitellus 1.50-1.8 mm., av. 
i 65, inner envelope 3-4 mm , av. 3.5 mm., outer 
envelope 4.1-5.4 mm., av. 5 mm.?] 

gg. Eggs brown above and yellow below, inner envelope 
distinct, 13-3.0 mm, outer envelope 3.6-5.0 mm.; 
vitellus 1.6-1.9 mm. Egg complement 1000 to 3000. 
Season April 6 to May 18. Rana palustris 

Plates VIII-4; 1X^47 



GENERAL ACCOUNT 43 

ff. Egg mass a plinth, complement large, 1000-10,000 eggs; eggs black above 
and white below. 

g. Outer envelope large, 10-15 nim., inner envelope indistinct, some- 
times absent, 5.0 6.0 mm., vitellus x-x.8 mm., black and white, far 
apart in mass like R. syhatica; egg complement 1000-1000 eggs in 
masses as large as 8 X 6 in Northwest. Season March to May. 

Rana p. prctiota 

gg Outer envelope smaller, 7 o mm. or less, inner envelope i 4-4.0 mm., 
vitelli i.4~x.5 mm , black and white, close together like R p/p/wr 
masses Except for R. p/p/cns, east of Rockies, 
h. Vitellus av xa-x.4orx5mm 

i. Vitellus av. x 5 (range x 4 x 6 mm.). Outer envelope 4.5-5.0 
mm , distinct, inner envelope 3.x mm. Masses larger, 5000 
10,000 eggs Midwest. Season March to May 15. 

Rana arcolata circulosa 

li. Vitellus av. x o mm (range 18x4 mm ), inner envelope 3.1 
4 4 mm , outer envelope 4460 mm , mode 5 x mm., av. 5.3 
mm. Egg complement 5000 or more Southeast. Season March 
13 to Nov 3 Ram captto 

Plates VIII x, IX 44, XV i, x, XCII 4 

hh. Vitellus av 1.4 i 7 mm. (range 13x0 mm ), inner envelope x 3 - 
3 x mm , outer envelope 3466 mm. Egg complement 1000 5000. 
i. Av. outer envelope 5 o mm or larger. 

) Av. outer envelope 6 3 mm. (5666 mm ), inner envelope 
x 4 3 o mm , av. x 75 mm., vitellus j 4 av., range 1.3 i 6 
mm , vitellus black or brown above, white or yellow below 
Egg complement 800 1500. Season June X5 to July 30 

Rana tcpttntnonahs 
Plate CXVIII 3 

jj. Av. outer envelope 5 i mm (range 4x 6.0 mm., mode 5.0 
mm ), inner envelope 1534 mm., av. x X5 mm , vitellus 
j 7 mm., range i 3 x.o mm. Egg complement 3500 4500. 
Season March 30 to May 15 Rana p. piptws 

Plate 1X^48,0-3, 4 

jjj Av. outer envelope 3 8 mm (range 3454 mm , mode 4 o 
mm ), inner envelope x 4 3 x mm , vitellus i 4 1.8 mm., 
av. 1.6 mm., egg complement 1084. Season January to De- 
cember. Rana p. sphcnoccphala 

Plate IX-49 



44 HANDBOOK OF FROGS AND TOADS 

cc. Three envelopes. 

f. Outer envelope 3.8-4.5 mm., av. 40 mm.; firm; middle envelope 
1.6-3.4 mm., av. 1.8 mm.; inner envelope 1.3-3.0 mm., av. 1.5 mm. 
All envelopes distinct. Vitellus 1.9-1.5 mm., av. 1.1 mm., black and 
white. Mass compact i X 2. X 1.1 in. to i X 4 X M in. Eggs 
firmly attached to each other. Egg complement 900-1100. Season 
March I to May i. Rana b. boylti 

Plate VIII -5 



ff. Outer envelope 6.4-14.0 mm. 

g. Egg mass small, 400-500 or less, all envelopes distinct. 

h. Outer envelope 6.4-7.9 mm., av 7.1 mm.; middle envelope 
4.1-5.0 mm., av. 4.6 mm., inner envelope 1.75-4.8 mm., av. 
3.9 mm., vitellus 1.8-1.3 mm., av. 1.1 mm., black above and 
light gray tan below, phnthhke mass 18 X 40 mm. with eggs 
J^-J^ in. apart. Egg complement 100-400 eggs. High Sierras. 
Season June, July. Rana b. stcrrae 

hh. Outer envelope 1.1-1.1 mm., middle envelope 5-6 mm., inner 
envelope 4.9 mm., vitellus 1.15 mm , black above, cream be- 
low. Mass 400-415 eggs. Washington and northern Oregon. 
Season last of May to July i. Rana cascadae 

gg. Egg mass large, 750-4000. Envelopes quite distinct or indistinct, 
h. All envelopes indistinct, especially the middle one. Outer 
envelope 10-14.0 mm., middle envelope 6.1-8.0 mm., av. 6.8 
mm., inner envelope 40-6.7 mm., av. 5.7 mm , vitellus 1.3- 
3.6 mm., av. 3.0 mm , black above, creamy white below. 
750-1400 eggs in a flat mass 6 X 10 in. across, vitelh * in. 
apart. Principally northwest Oregon northward. Season Feb- 
ruary to May. Rana a. aurora 

hh. All envelopes quite distinct, middle one particularly dense 
and distinct, outer envelope 7.5-11.8 mm., av. 8.5 mm., mid- 
dle envelope 3.9 6.4 mm., av. 4.40 mm., inner envelope 3.9-- 
6.4 mm., av. 4.4 mm. Vitellus 1.0-1.8 mm., av. 1.1 mm.; 
black and creamy white, egg complement 1000-4000 eggs 
in a soft, viscid mass 1.6 X 4 X 3 inches to 6 X 4 X 4 in. 
Principally California and Lower California. Season January 
to March. Rana a. draytonri 



GENERAL ACCOUNT 45 

cc. Egg mass a loose irregular cluster, sediment-covered. 

d. Egg mass small, generally about or less than i in. (1.5-3.5 cm O m 
diameter, normally 10-15 or 4 r * r ely 100 or more in one mass, 
e. No inner envelope. 

f. Smaller species 19-37.5 mm. (19-31 males, 19-37.5 females). 
Egg or vitellus 0.615-1.1 mm., av. below 1.1 mm. (except in 
P. . septentrtonalts, 1.1-1.4 mm.) single envelope 1.6-5.0 mm. 
normally, rarely to 7.8 mm. 

g. Vitelh 1.1-1.4, envelope larger. Season May and early June. 

Pseudacris nigrita septentrionalts 

gg. Vitelh 0.615-1.1 mm.; envelope 1.6-5.0 mm. normally. 
Season early February or earlier to May. 
h. Envelope 1.1 1.4 mm. or more, vitellus 0.615- 9 mm. 
Season March 10 to May 10. Pseudacris n. darkii 

Plates VII-4, L~4 

hh. Envelope i 6-1.8, vitellus 0.9 i.o mm. Florida to Georgia. 
Season February to August 15. Pseudacrts ;/. verrucosa 

hhh. Envelope 3.0-5.0, sometimes more. 

i. Envelope 3 1-4.0 mm., vitellus 0911 mm. Vitellus 
black and cream or slightly yellowish. Season February 
to May. Pseudacrts n. fertarum 

Plates VII 1,5, LI 4 

ii. Envelope 3.0- 5.9, sometimes 7 8 mm., vitellus 0.9 1.1 

mm. Egg complement 500 800 Season March 19 to 

May. Pseudacrts n. truer tat a 

Plates VII-6, IX } ia,b,c, LIII 4 

ff. Larger species 14 48 mm. (14 41 males, 17-48 mm. females). 

Egg or vitellus av. over i i mm. (range 1.1-2.0 mm ). Single 

envelope 3.0-8 5 mm. Masses IQ 100, rarely 100 in a mass. 

g. Envelope 6.0 8.5 mm., vitellus 1.6 mm. West Virginia to 

Ohio to northern Mississippi and Alabama. Season March 

to July. Pseudacris brachyphona 

gg. Envelope 3.0-7.15 mm., av. normally under 6.0 mm. 

h. Vitelh 1.6 mm. Envelope 3.1-4.1 mm., av. under 4.0 mm. 
Southeast. Season November to March. Pseudacris ornata 



46 HANDBOOK OF FROGS AND TOADS 

hh. Vitclh i.z-i.3 mm. (Bragg; 1.1-1.8 or 10, Strecker's ma- 
terial), envelope 3.0-7.15 mm , av. over 5.0 mm. Texas to 
Oklahoma. Season December to late May. 

Pstudacris strtckcri 
ee. Inner envelope i 4 1.7 mm. 

f. Outer envelope smaller, 1.1-3.0 mm., vitellus smaller, 0.65-1.1 
mm., inner envelope 1.4-1.1 mm. 

g. Outer envelope small, 1.1-1 4 mm. or more; inner envelope 
1.4 1.8 mm., vitellus 0.65-0.9 mm , mass smaller, 13-30 eggs; 
egg complement, 150-300 eggs. Season March 5 to mid-August. 

Pscudacns n. clarkti 
Plate VII-4, L- 4 

gg. Outer envelope larger, 3.0-5 o or 7.8 mm., inner envelope 1.1 
mm.; vitellus o 91 i mm Mass 15-300 eggs per mass. Egg 
complement 500 -1500 eggs. Season March 10 to May 10. 

Pseudacrn n. tnsmata 
Plates VII-6, IX 3 ia,b,c, LIII-4 

ff Outer envelope larger 4.7-6 7 mm., inner envelope i 8-1.7 mm.; 
vitellus larger 13135 mm , vitellus light brown above and pale 
white, yellowish, or greenish below. Masses 9-75 eggs, rarely 
single. Egg complement 600- 1500. Hyla regilla 

dd. Egg mass an irregular cylinder 1-6 in (i 5-15 cm.) in length, extending 
along plant stem or grass blade, envelope single, 3.8-5 6 mm , vitellus 
i .4 -i.o mm. Egg complement 1331. Season April or earlier to August 17. 
(See also d, dd, c, cc under c Eggs laid in bands which soon become 
loose cylinders extending along plant stems or grass blades ) 

Scapbtopus fy. bolbrooktt 
Plates -1,4, IX- 5 ,XXI- 5 

We never described in detail our results from work in Texas during 1925- 
19^5. The Florida group of Carr, Gom, and Van Hynmg have added much 
to our knowledge of the Florida forms. H. M. Smith in 1934 did a good study 
of the Kansas forms. J. R. Slater of Tacoma has described the forms of the 
Northwest (still in manuscript), and A. and R. D. Svihla and K. Gordon 
have also added data about that area. But the group which has been most 
vigorous is the Oklahoma group (from the University of Oklahoma Bragg, 
Trowbridge, Ortenburger, and others; and from Oklahoma A. and M. Col- 
lege, the Moore group). 

Tadpoles (Plates X-XV: lateral views, X-XI; mouth parts, XII-XIV; de- 
velopment, XV) : Some 25 to 30 species of tadpoles of the United States and 



GENERAL ACCOUNT 47 

Canada need to be described. Most have been found but were not described, 
and the tadpoles of 15 forms are yet unknown to science. The life histories 
of the robber frogs we know in our country indicate no free tadpole stage, 
the whole development being in the egg. 

In discussions of the size of tadpoles quite small is i inch (24 mm.) or 
smaller; small, T-I% inches (24-3*5 mm.) : medium, i 3 -2 inches (40-50 mm.) ; 
large 2%-$% inches (60-86 mm.); quite large, ?/. : >-4 inches (95-100 mm.); 
very large, 5%-5 Jfi inches (135-145 mm.). Some of the toads and swamp 
cricket frogs may have tadpoles a little less than i inch (23 or 24 mm.) in 
length, whereas bullfrogs may have tadpoles 5% inches (145 mm.) or even 
larger. 

The tadpole has a body and a tail. The body has sensory lines, a breathing 
pore or spiracle, a vent or anus, eyes, nostrils, and a mouth. The mouth is 
wholly unlike the adult mouth. It usually has a disk called a Libium (with 
upper and lower labia). Generally about the edge of the labium are tubercles 
or papillae. At the inner edge of each labium or at the very portal of the mouth 
opening itself are horny crescents called upper and lower mandibles. On the 
upper and lower labia arc horny ridges of teeth or combs for scraping food. 
The tail has two parts: the axis consists of muscle segments; and the fin con- 
sists of upper and lower crests. 

The narrow-mouthed toad tadpole has no mouth disk, no labial teeth, no 
papillae, no horny beak. The spiracle is next to the vent. The eyes are on a 
lateral ndgc. These tadpoles are small black flattened creatures with some 
white on the tail axis or body. 

The ribbed toad (Ascaphus) has a mouth disk, an upper mandible only, 
papillae on the lower lip or labium, upper labial teeth at least two rows to a 
ridge, labial tooth ridges two to three above and seven to ten below, a spiracle 
in the middle of the venter nearer the hind legs than the snout, and eyes 
straight back of and close to nostrils, dorsal, and equidistant from micldorsal 
line and lateral outline when viewed from above. The tail is spatulatc and 
rounded as in some mountain stream tadpoles. The tadpoles are black or 
brown with white tips. 

The spadefoots have the vent in the middle position, the spiracle below the 
body axis and on the left side, the papillae completely around the labium 
except for a small interval above (absent in one species), the papillary border 
not emargmate on the side, labial teeth three to six ndges above and four to 
six ridges below, eyes dorsal and nearer middorsal line than lateral outline, 
and muscle segments of the tail plainly visible. 

The toads usually have small blackish tadpoles with anus median, papillae 
confined to the sides of the labium, upper and lower edges of the labium 
toothed, sides of labium emarginate, labial teeth two ridges above and three 
ridges below, and eyes dorsal and slightly nearer the lateral outline than mid- 
dorsal line. The spiracle is on the left side and is small. 







;;;,;:, :;; 




Plate X. Mature tadpoles (X 1 )- i- Anderson tree frog, Hyla andcrsomt, 2. Green tree frog, Hyla 
cinerca. 3. Peeper, Hyla cmctfct cntcifer. 4. Pine woods tree frog, Hyla jcmonihs. 5,6. Barker, 
Hyla gratiosa. 7. Squirrel tree frog, Hyla Stjtwella. 8. Little chorus frog, Pscudacns octdans. 9,10. 
Common tree toad, Hyla vcrsicolor vcrsicohr. u. Cricket frog, Acns gryllns gryllus. 12. Spadefoot, 
St'aphiopus holbrookti holbroofyi. 13. Oak toad, Bttfo qttercictts. 14. Narrow-mouth toad, Micro- 
hyla carolincnsis. (Wright, Gen., 1932, pp. 46-47.) 



;,^w:^ vf.' 






13 




X/. /? tadpoles (mature XO- 1 2 - Gopher frog, /?. r0/7/'/0. 3,4. Green frog, /^. damitans. 
5,6. Southern bullfrog, K. grylto. 7,8. River-swamp frog, /?. hec^schcn. 9,10. Southern meadow frog, 
R. piptcns sphenocephala 11. Mink frog, R. septentnonahs. 12. Meadow frog, R. p. pipiens. 13. 
Sphagnum frog, R. virgatipcs. 14. Pickerel frog, R. palustris. (Wright, Gen., 1932, pp. 45-47.) 




rlatc XII. Tadpole mouth parts (Hyla, Acris, Pseud acris, Bttfo). i. Peeper, H. crucijer crucijer. 
2. Barker, H. gratiosa. 3. Anderson tree frog, H. andersonn. 4. Green tree frog, H. cinerea. 5. Eastern 
swamp cricket frog, P. nigrita jcnarum. 6. Cricket frog, A. gryttus gryllus. 7. Fowler's toad, B. 
woodhousii jotilcn. 8. Little chorus frog, /'. octtlans. 9. Squirrel tree frog, H. sqmrella. 10. American 
toad, B. amencamts americanus. n. Southern toad, B. tetrestris. 12. Common tree toad, H. vcrsicolor 
versicolor. 13. Pine woods tree frog, H. fcmorahs. (Wright, Gen., 1932, pp. 54"55-) 




Plate Kill Tadpole mouth parts (Rana, Hyla, Pscudacrts, Bu)o, Ascapftus). i. spnagnum irog, 
R. virgatipes. 2. Nevada frog, R. fisheri. 3. Canyon tree frog, H. arenicohr. 4. Clarke's chorus frog, 
P nignta claret. 5. Spotted toad, B. punctatus. 6. Spadefoot toad, B. compactihs. 7. Nehulous toad, 
B. valliccps. (Wright and Wright, MS, 1930.) 8. American bell toad, A. truci. (After Gaige,) 



52 HANDBOOK OF FROGS AND TOADS 

The frogs have medium-to-large tadpoles, with spiracle on left side, vent on 
right side of lower tail fin's base. The spiracle is near the body axis; the papil- 
lary border is cmarginatc on the side; the labial teeth are two or three ridges 
or more above and three or four or more below. Some may have spatulate 
tails, if high mountain forms, others may have upper tail crest high and far 
on the body like a tree frog tadpole, but most have the tail crests neither very 
high nor extending far on the body. 

The tree frogs have small or medium tadpoles, often with high dorsal tail 
crest extending far on the body, the vent on right side of ventral tail crest, and 
the spiracle on left side near the body axis. The labial teeth have two ridges 
above and two or three below. 

The following synopsis of tadpoles is offered only after considerable de- 
bate. We could have put it in the first edition. No one knows the extent of 
variation in the mouth-part characters. If one attempts to use other workers' 
results, he encounters personal equational differences in measuring technique. 
We might find no median space in the second upper lateral tooth row in 
some P. n. clarion and a median space in others. Dr. Bragg figures his material 
with no median space, as we at times have found. What is normal for the 
whole range of a species? This synopsis is presented in the hope that some- 
one will attempt to learn the frog tadpoles of the United States in the field 
and refine our descriptions. Grace L. Orton may. 




Plate XIV. Tadpole mouth parts (Rana, Mtcrohyla, Scaphiopus). i. River-swamp frog, R. 
hecfahen. 2. Young of (i). 3. Mink frog, R. scptcntrwnahs. 4. Sphagnum frog. R. virgatipcs. 5. 
Southern bullfrog, R. grylto. 6. Narrow-mouth toad, M. carohncnsis. 7. Green frog, R cl-"""" f 
8. Spadefoot, S. holbrootyt. (Wright, Gen., 1932, pp. 54-55.) 



4 HANDBOOK OF FROGS AND TOADS 

SYNOPSIS OF UNITED STATES TADPOLES 

A. No tadpole stage. 

Eleutherodactylus ricordn plantrostns (Plate LXXVIII-5,7,8), E. latrans; 
Syrrhophus camp; S. gatgeae; S. marnockii. 
AA. A tadpole stage in water. 
B. Tadpoles unknown. 

Scaphiopus holbrookii albus; Bufo caltfornicus; B. hemiophrys; Pseudacris 
n. nigrtta; P. occidental**; P. n. septentrional ts; Hyla baudtnii; H. v. 
phatocrypta; H. avwoca; H. vcrsicolor chrysosccltsj Rana a. aurora; R. 
prettosa; Mtcrohyla anolata; Hypopachus cuneus 14 in all. 
BB. Tadpoles seen but not described in detail. 

Bufo alvarius; B. b. boreas; B. canorusj B debths; B. woodhousii fowleri; 
Pseudacris ornata; Hyla cinerea cvittata; H. wrtghtorum; Rana a. areolata; 
R. sylvattca cantabrtgensis; R. fisher*. 
BBB. Tadpoles in this synopsis. 

a. Mouth disk absent, no labial teeth; no papillae, no horny beaks; 
spiracle median near anus; nostril within edge of mouth fold, eye on a 
canthus; tadpoles (2.3-16 4 mm.) small, black or grayish olive tadpoles 
with a stripe on the middle of the tail musculature. 

BREVICIPITIDAE 

b. Tail tip always black, eyes just visible from ventral aspect; black 
pointed excrescences back of upper labial edge, lower mandibular 
prolongation gray, coloration citrine-drab or grayish olive, with 
middorsum dark grayish olive, venter with white or pale pinkish 
cinnamon spots, sides of body without striking longitudinal light 
bands, light band at base of tail musculature not prominent (in 
alcohol). Outer egg envelope not truncate, mass seldom showing 
distinct outline of each egg envelope in a mosaic fashion. Texas 
westward to Fort Davis Mountains. Mtcrohyla olwacea 

Plate CXXVI-4,5 

bb. Tail, sometimes with black tip, eyes plainly visible from ventral 
aspect, inner face of upper labial edge with no black pointed ex- 
crescences; lower mandibular prolongation light, general coloration 
black with purplish gray or hair brown dots, venter with white 
or yellowish bands and large blotches, sides of body with same 
coloration; light band at base of tail musculature prominent (even 
in alcohol). Egg with truncate outer envelope, giving mass a mosaic 
appearance on water's surface. Virginia to Florida to Texas and up 
Mississippi River to Indiana. Mtcrohyla caroltnensts 

Plates X-i 4 , XIV-6; CXXV- 4 



GENERAL ACCOUNT 55 

aa. Mouth disk present; upper and lower labial teeth, labial maxillae; at 
least the upper horny beak present, nostril free of mouth, 
b. Spiracle median nearer insertion of hind legs than tip of snout; labial 
teeth 1/7 to 3/10, upper labial teeth at least two rows to a ridge; no 
lower beak; many rows of papillae on edge of lower labium, mouth 
large and round; anus median, black or blackish brown tadpoles 
speckled with black; tail may be of body color or spotted with 
creamy white, tail tip white or yellowish white; upper tail crest not 
extending on the body. Washington to California. 

Ascaphus truet ASCAPHIDAE 
Plates XIII-8, XVI-i 

bb. Spiracle sinistral, upper labial teeth not with two close rows on each 
ridge, labial teeth 1/3, 1/1, 1/3, 1/4, 3/3, 3/4, 4/4 to 6/5, upper 
and lower horny beaks; papillae on lower edge of labium absent or 
in one or two rows. 

c. Anus median, spiracle lateral below the lateral axis (of tail mus- 
culature, projecting) sometimes as much ventral as lateral, upper 
tail crest extends on the body to a vertical nearer hind legs than 
the spiracle or only halfway; viscera visible (in preserved speci- 
mens) through the skin of the belly. Eyes distinctly dorsal, 
d. Labial teeth 3/4 to 6/6, papillae extending completely around 
the border of the labium except for a short-toothed median 
interval above (sometimes absent in one species); papillary 
border not emarginate on each side, eyes nearer middorsal line 
than lateral outline, on lateral axis, tadpoles 14.5-65 mm. in 
length, spiracle a slit, very low on side, about on the level 
of the mouth, in general very bronzy tadpoles, myotomes of 
tail musculature well indicated. Tail tip usually rounded. 

SCAPHIOPODIDAE 

e. Teeth 6/6, 5/6, 6/5, 5/5, tadpoles small, 2.1-18 mm., trans- 
forming at 8.5-11.0 mm., inner papillae present, spiracle 
equidistant between eye and base of hind legs or vent, eye 
1.4-1 8 nearer tip of snout to the spiracle, av. 1.51, inter- 
nasal space in interorbital space 1.18-1.83, av - I -5^J depth 
of body 1.75-1.5 in body length, av. 1.04, muscular part 
of tail in depth of tail 1.45-1.5, av. 1.98, last lower row 
of teeth longer than horny beak. Last lower row of teeth 
1.5 times in next to lowest row of teeth. Egg mass an ir- 
regular cylinder, at first bandhke. Massachusetts to Florida 
to Texas and Arkansas. Scaphiopus h. holbrookii 

Plates X -ii, XIV-8 



5 6 HANDBOOK OF FROGS AND TOADS 

cc Labial teeth normally 4/5 or 4/4, sometimes 5/4, 3/4, or possibly 1/4; 

inner papillae generally absent, tadpoles small or large; eye 1.0-1.6 

nearer tip of snout than spiracle; last lower row of teeth less than or 

J^ of the width of horny beak. 

f. Tadpole large (65 mm ), transforming at 13 -2.4 mm.; teeth 4/4, rarely 
5/4 or 3/4, upper fringe of papillae broken in middle by a row of teeth; 
eye 1.1-1.3 nearer tip of snout than to the spiracle, av. i i, muscular 
part of tail 1.15- i 66 in depth of tail, av. 1.43 , width of body in its own 
length 1.55 -1.9, av. 1.66, depth of body 1.05-1.15 in body width, av. 
1.09, spiracle j 15 -i 6 nearer eye than base of hind legs or vent, av. 
1.37, internasal space 1.5-175 in interorbital space, av. 1.6, third 
lower labial row broken in middle, median interval of second lower 
row broad, last row of teeth 1.75-10 times in next to last row of 
teeth and usually i .7-1.0 times in mandibles, second upper row usually 
broken in middle. Egg mass a loose cylinder, many eggs on stalks. 
Montana to Texas and westward to Pacific Coast States. 

Scaphtopus hammondii 

(g. Upper labial mandible with beak, lower labial mandible with 
notch. Scaphtopus h. hammondii 

Plate XVlII-3,5 

gg Upper labial mandible without beak, lower labial mandible with- 
out notch. Scaphtopus h. bombifront 
After Bragg) 

[ggg. Probably belongs with gg. Scaphtopus h. tnttrmontanut] 

ff. Mature tadpole small (2.1-2.7 mm.), transforming at 7 5-11.5 mm., 
upper fringe of papillae is conspicuously broken in middle by a row 
of teeth or not so broken and the first row absent, last lower row of 
teeth 1.1-1 4 in width of mandibles. 

g. Teeth usually 4/5, sometimes 3/4, interval between lateral parts 
of third upper row i 5 -3 o in either lateral row or 5.0 in width 
of mandibles, interval between lateral portions of fourth row 5 o 
times in width of mandibles, second upper row normally broken. 
Eggs tandcmhke, or Bufo-likc or lattice work. (Compiled after 
Bragg Oklahoma and Arkansas to Texas. Scaphiopus h. hurteni 

gg. Teeth usually 4/4, rarely 5/4, 3/4, or possibly 1/4, interval be- 
tween lateral parts of third upper row equal to or 1/3 either lateral 
portion of this row or 1.5 times in mandible, interval between 
lateral portions of fourth upper row 1.3 in width of mandibles; 
second upper row normally unbroken, eye i. 0-1.6 nearer tip of 



GENERAL ACCOUNT 



57 



snout than to the spiracle, average 1.3, muscular part of tail 
1.1-3.33 ' m depth of tail, av. 1.7; width of body in its own 
length 1.3-1.6, av. 1.4, depth of body 1.15-1.55 in body width, 
av. 1.35; spiracle 1.5-1.1 nearer eye than base of hind legs or 
vent, av. 1.85, mternasal space 1.45-1.3 in mterorbital space, 
av. i.o, third lower labial row continuous; median interval of 
second lower row very narrow. Last lower row 1.5-3.0 
times in next to lower row of teeth; second upper row not or 
barely broken in middle, egg mass an irregular cylinder, at first 
bandlike. Texas to California and Mexico. Scaphtopus coucbii 

dd. Labial teeth 1/3; papillae confined to the sides of the labium (or lower 
half only in Bufo punctatus), upper and lower edges toothed, papillar 
border on each side emarginate, eyes slightly nearer lateral outline than 
middorsal line, above lateral axis, spiracle small, a porehke opening, 
e. Tadpoles 14-18 mm. in length (a few forms 35-56 mm ) 

BUFONIDAE 

f. Papillae only on lower half of lateral margin, or a slight marginal 
row of 4 to 6 papillae on upper half, no inner papillae normally, 
third lower row of teeth equal to the first row of lower labial 
teeth, median space between lateral parts of the second row of 
upper labial teeth 1-3 times in either lateral row, horny beak 
1.1-1.5 in third lower labial row, tadpoles to 15 mm One of the 
blackest Bufo tadpoles, tail musculature evenly dotted with black, 
venter with light grayish vmaceous spots, eggs single or film or 
scattered mass on bottom, not in files. Central Texas to California, 
Utah, and Lower California. Bufo punctatus 

Plate XIII 5 

ff. Papillae on upper and lower halves of lateral labial margin, some 
inner papillae. 

g. Bicolor tadpoles distinctly lighter below, not black tadpoles, 
upper tail fin highly arched, horny beak 1.3-1.5 in first or 
second lower row of teeth. 

h. Light grayish olive, dark olive-buff or clay color, the 
lightest of our toad tadpoles, in life intestine does not show 
through the skin of the belly (shows in preserved speci- 
mens), third lower row of labial teeth short, 1.8-1.4 in 
first lower row, third lower row 1.5-1 o times in the horny 
beak or 1.6-1.8 m longest lower row, median space between 
lateral parts of the second row of upper labial teeth large, 
1. 1-1.0 times greater than either lateral part, depth of tail 
in length of tail 3.0-3.5; horny beak 1.3-1.4 in first or 



58 HANDBOOK OF FROGS AND TOADS 

second row of teeth; lower loop of papillae to end of third lower but 
not under it, usually not two rows; tail musculature with dark 
vinaceous drab band to tail tip; below this a pale vinaceous pink 
band, belly pale vinaceous pink; eggs in files or strings, brown or 
buffy brown, cream cr straw yellow below; no inner tube. Texas to 
Mexico, Arizona. Bufo compactilis 

Plate XIII-6 

(hh. Doubtless Bragg's (1936) Bufo cognatus belongs in g. Dorsal surface 
mottled brown and gray silvery areas interwoven with black ones. 
The ventral surface markedly lighter than the dorsal. The viscera 
(in mature tsdfcles) cnly slightly or not at all visible, lower tail fin, 
clear or almost so, third lower rcw of teeth 1.5-1 8 times in horny 
beak, or 1.5-3.0 in longest lower row, median space in second upper 
row (none says Bragg, 1.0-1.2. in either lateral part says Smith), 
depth of tail in length of tail 1.5, eggs in single row sometimes 
double, an inner gelatinous envelope and an outer tougher envelope 
or tube, somewhat scalloped in outward appearance. (After A. N. 
Bragg, Okla., 1936, p. 19.) Minnesota to Texas and southern Cali- 
fornia. Bufo cognatus) 

gg. Elack or blackish tadpoles, in life the intestine shows through the skin 

of the belly, third lower rcw of labial teeth long, 1.1-1.6 in first lower 

row, third lower row 1.0-1.5 in horny beak; median space between 

lateral parts of the second row of upper labial teeth small, contained 

1.0-4.9 times in either lateral half, not greater than lateral half. 

h. Papillae very faint, minute, at times hard to see but present; third 

lower rcw of labial teeth not equal to first lower rcw of teeth, but 

equal to or greater than horny beak, horny beak 1.1-1.5 in upper 

fringe, 1.3-1.5 in first lower row; median space of second upper row 

1-4 times in either lateral part of it; upper edge of tail musculature with 

8- 10 black bars with intervening pale olive-buff areas; irregular black or 

brown band on tail with cartridge buff or tilleul buff below it; tadpoles to 

2^ mm. Eggs in files, inner tube present, one or two rows of eggs. 

Louisiana to Costa Rica. Bufo valliceps 

Plate XIII-7 

hh. Papillae plainly visible. 

i. Tadpoles of Rockies to eastward. Tadpoles usually less than 30 

mm., third lower row of labial teeth 1.1-1.5 * n fi fst row - 

j. Tadpole to 14 mm.; horny beak in upper fringe 1.75, in first or 

second lower row 1.5 , horny beak equal to or less than the third 

lower row of teeth; median space between two parts of second 



GENERAL ACCOUNT 



59 



upper row 1.4-1.1 in cither lateral part; third lower row of teeth in first 
lower row i .1-1.4; two or more rows of strong papillae from end of upper 
fringe to end of third lower row, sometimes 3-5 rows at side of labium; 
lower loop of papillae far below level of third row and with at least two 
rows of papillae, mouth in interorbital space 1.0-1.5, av - I - I 7> mouth 
larger than internasal space 1.1-1.6, av. 1.36; depth of tail in tail length 
1.4-4.5, av. 3.17, eye nearer snout than spiracle 1.0-1.57, av. 1.1; nostril 
nearer eye than snout 1.1-1.8, av. 1.48. Egg mass long file, inner tube 
with no partitions. North Carolina to Florida to Louisiana. 

Bufo temstris 
Plate XII -ii 

jj. Tadpole to 17 mm.; horny beak in upper fringe 1.1-1.5, * n ^ rst or second 
lower row of teeth 1.1-1.1 or 1.1-1.3; horny beak greater than third row 
of lower labial teeth; median space 1.15-1.0, 1.0-4 > or I -3~3- * n either 
lateral part, third lower row of teeth 1.3-1.5 or 1.4-1.5 in first lower 
row; one row of weak papillae from upper fringe to end of third lower 
row of teeth with a few scattering papillae at the side of the labium, 
lower loop with only two or three scattering papillae beside the outer 
row of weak papillae. 

k. Mouth in interorbital distance 0.77-1.0, av. 0.91; horny beak in upper 
fringe 1.1-1.4, horny beak in first or second row 1.1-1.1 times; third 
row in first lower row 1.3-1.5, depth of tail in tail length 1.15-1.7, 
av. 1.97, spiracle nearer eye than vent 1.04-1.54, av. i 18; eye nearer 
snout than spiracle i 0-1.17, av. 1.16, mouth larger than internasal 
space 1.4-1 i, av. i 76; papillae of lower labial loop do not extend 
under the end of the third row of labial teeth, tail musculature in tail 
depth 1.16-1.66, av. 1.04, internasal space 1.1-1.8 in interorbital dis- 
tance, av. 1.6, spiracle 1.05-1.55 nearer eye than vent, av. 1.18. Egg 
mass long file, partitions, inner tube present. Eastern North America 
from Hudson Bay southeast. Bufo a. amcrtcanus 

Plate XII-io 

kk. Mouth in interorbital distance 0.9-1.5, horny beak 1.4-1.9 in first 
upper fringe of teeth, horny beak in first or second row 1.1-1.4; 
depth of tail in tail length 1.75-3.8. 

1. Mouth in interorbital distance 1.07-1.5, horny beak in upper fringe 
1.4-1.5, horny beak in first or second row 1.1-1.3, third lower row 
1.3-1.6 in first lower row; depth of tail in tail length 1.88-3.83, 
av. 3.33, spiracle nearer eye than vent 1.15-1.7, av. 1.45, eye nearer 
snout than spiracle 1.0-1.6, av. 1.16; mouth larger than internasal 
space i . i-i . 83 , av. i .47, papillae of lower labial loop extend slightly 
or not at all under the end of the third lower row, tail musculature 
in tail depth 1.6-1.3, av. 1.85; internasal space 1.5-1.16 in inter- 



60 HANDBOOK OF FROGS AND TOADS 

orbital space, av. 1.86; spiracle 1.15-1.7 nearer eye than vent or 
base of hind legs, av. 1.45. Egg mass long file, no inner tube, 
sometimes two rows of eggs. Massachusetts to Missouri on the 
north and South Carolina to Texas on the south. Bufo w. fowleri 

Plate XII-7 

11. Horny beak in upper fringe 1.8-1.9; in first or second lower row 
of teeth 1.6; third lower row 1.3-1.4 in first lower row, depth 
of tail in length of tail 1.75; spiracle nearer eye than vent 1.15; 
eye nearer snout than spiracle i.o-i.i or equidistant, papillae of 
lower labial loop slightly extends under the ends of the third 
lower row; tail musculature in tail depth 1.1-1.5; egg mass a 
long file, no inner tube, sometimes two rows of eggs. (After K. 
A. Youngstrom and H. M. Smith, Kan., 1936, p. 633.) Western 
Iowa south through Texas into Mexico and west to Idaho and 
Imperial Valley, California. Bufo w. woodbousii 

ii. Tadpoles of Pacific Coast area. Tadpoles often over 30 mm. (maximum 
17-56 mm , 2.7 canorus, 33 exsul, 19 nelson^ 44 borcas, 56 b. halophtlus 
boreas group). Third lower row of labial teeth 1.4 or 1.5-1.0 in first lower 
row. 

j. Smaller tadpoles (17 mm. canorus, 35 mm. exsuf). Median space in 
second upper row of teeth almost equal to or 1.1 times in either lateral 
portion of this row, third lower row 1.5-1.0 in first and second lower 
rows, 1.1-1.5 * n f irst upper fringe. 

k. Third lower row 1.5 in first lower row, i.o in second lower row or 
1.1-1.3 i fl upper fringe. (After Storer.) Yosetnite region. 

Bufo canorus 

kk. Third lower row 1.4 in first or second lower row or equal to or 
1.5 in first upper fringe. Deep Springs, Calif. Bufo exsul 

}]. Larger tadpoles (19 mm. nelsoni, 55 mm. b. halophtlus}. Median space 
in second upper row of teeth 1.0-3.7 in either lateral rows, third lower 
row 1.4-1.0 in first or second lower row. 

k. Median space in second upper row 1.0^-1.5 times in either lateral 
portion of this row or 1.5-3.0 in horny beak; third lower row of 
teeth i times in first or second lower row of teeth. Southern Nevada 
into California. Bufo b. nelsoni 

kk. Median space in second upper row of teeth 3-2.-3.7 in either lateral 
row or 3.8 in horny beak; third lower row 1.4 in first or second 
lower row or 1.5-1.6 in first upper fringe. (After T. I, Storer.) 



GENERAL ACCOUNT 61 

Montana and Colorado to northern California 
and British Columbia, possibly Alaska. 

Bufo b. halophtlus 

[The characters used for this boreas group are 
trivial and not to be given too much significance.] 

ee. Tadpoles 55 or 56 mm. (body 2.3 and tail 33; see Storer for descrip- 
tion). Bufo b. balophtlus 

Anus dextral, spiracle distinctly lateral on or near body axis. 

d. Papillary border on side with an emargination, tadpoles 50-149 mm. 
in length; papillary fringe on upper labium extends not at all inward 
beyond the end of the upper fringe of teeth or only 1/8 to 1/16 of the 
length of the fringe; length of horny beak in upper fringe of teeth 
i.o^-i 5 times; labial teeth 1/3 or 3/3 or 3/4, rarely to 7/6. 

RANIDAE 

e. Labial teeth 7/6 to 3/3, not customarily 1/3, tadpoles 47-88 mm. 

f. Labial teeth 7/6 or 6/6, depth of tail in length often 3.3-4.1. 

(See T. I. Storer for description ) Coastal California and Oregon. 

Rana b. boy It i 
ff. Labial teeth 1/4, 3/4, 4/3, or 5/3, not 1/3. 

[R. aurora aurora, labial teeth 3/4, belongs here; no detailed de- 
scription on record, probably some R. a. draytom also fall here 
though some are 1/3 (T. I. Storer).] 

g. Labial teeth 5/3 or 4/3, tadpoles large to 88 mm., tadpole 
tail very prominently spotted and mottled, all three lower 
labial rows long, median space of second upper row 1.3-1.5 
in either lateral section of this row, tail tip pointed or 
rounded, depth of tail in tail length 3.3-3.4 times, spiracle 
1.1-1.3 ne &rer vent than tip of snout, nostril 1.3-1.8 nearer 
eye than snout, spiracle 1.8-1.0 nearer eye than vent. New 
Mexico, southern Arizona to central Mexico. 

Rana tarabumarat 
Plate CXXII-i,6 

gg. Labial teeth 3/4, tadpoles 49-71 mm. in length. Dorsal crest 

very high extending to vertical of the spiracle (Rana sylvattca) 

or tail broader nearer its tip than at its body insertion, 

rounded, spatulate, or elliptical (R. b. surrai); upper fringe 

of teeth about 1.5 times the length of the horny beak. 

h. Tadpole to 49 mm.; dorsal crest very high, extends on the 

body to vertical of the spiracle; tail tip acuminate, tail 

musculature begins to taper at once; dorsal crest higher 

in cephalic half; depth of tail in length of tail 1.9-3.1, 

av. 1.5; spiracle 1.5-1.0 nearer base of hind legs or vent 



62 HANDBOOK OF PROGS AND TOADS 

than tip of snout, av. 1.75; nostril equidistant from snout and 
eye; internasal space in interorbital space 1.7-1.9, av. 1.5; depth 
of body 1.08-1.13 in body width, av. 1.14; depth of body 1.66- 
z.o in body length, av. 1.78, second upper lateral row about 
1/5 of the upper fringe of teeth, the median space between the 
lateral portions of the second row 0.4-0.8 times either lateral 
portion; the third upper row is 1/4 to 1/9 of the upper fringe; 
fourth lower row of teeth is 6/n to 7/11 of the first lower row; 
spiracle 1.0-1.47 of the first lower row; spiracle 1.0-1.48 nearer 
eye (5.0-6.0 mm.) than base of hind legs or vent (4 5-7.6 mm.). 
Egg mass submerged, globular. Ontario to Nova Scotia, south 
to South Carolina, west to South Dakota and south to Arkansas. 

Rana s. sylvatica 

hh. Tadpole to 71 mm.; dorsal crest low extending on body to a 
vertical twice nearer hind legs than spiracle; tail tip rounded, 
elliptical, or spatulate; tail musculature for an inch or more 
does not taper; dorsal crest narrow in cephalic half, depth of 
tail in length of tail 1.5-5.7, av. 4.0; depth of body 1.1-1.8 in 
body width, av. 1.43, depth of body in body length 1.0-3.0, 
av. 1.315, spiracle 1.0-1.33 nearer base of hind legs or vent than 
tip of snout, av. 1.17; spiracle 1.45-1.375 nearer eye (4.0-8.0 
mm.) than base of hind legs or vent (5 8-n.o), av. 1.05; young 
tadpoles occasionally with teeth 1/4, rarely 3/3; nostril i.o- 
1.75 nearer the eye than snout, internasal space 1.3-1.0 in inter- 
orbital space, av. 1.5; second upper lateral row 1/4 to 1/9 of 
the upper fringe; median space between lateral portions of sec- 
ond upper row 1.0-1.3 times either lateral part; the third upper 
row is 1/9 to i/n of the upper fringe; the fourth lower row of 
teeth is 1/3 to 1/4 of the first lower row. The Sierras, California. 

Rana b. sicrrac 

ggg- Labial teeth 1/4; fourth lower labial row 1/4 of length of other 
three; median space in second row small. (After J. R. Slater, Wash.) 
Tadpoles 40-50 mm. Northern Oregon into Central Washington. 

Rana cascadae 
ee. Labial teeth 1/3, occasionally 3/3, rarely 1/3. 

(We have seen no tadpoles of R. prcttosa or R. p. lutcwentns in the field; 
we have seen preserved tadpoles with teeth 3/3. "In prcttosa there are 
four rows of labial teeth, one very short upper row and three lower rows. 
In luttiventns there are five rows, two long upper rows and three lower, 
as figured by Thompson (1913). The first lower row in lufetwnfris is longer 
than the analogous one in pretiosa" (A. Svihla, Wash., 1935, p. i").) 
Tadpoles 74-150 mm. in length; dorsal crest not extending to vertical of 
the spiracle but usually just ahead of the buds of the hind legs; tail 
always elliptical not spatulate; upper fringe of teeth equal to or slightly 
larger than (never 1.5 times) the horny beak. 



GENERAL ACCOUNT 63 

f. Tadpoles 74-84 mm.; tadpoles usually transform the same season they are 
born; transformation sizes 18-30 mm., av. 2.4 mm. (except R. arcolata 
group), tadpoles (except in R. capita or arcolata group) not strongly pig- 
mented on belly, viscera plainly or slightly showing through skin (in 
spirit specimens). Egg mass globular or plinthlike beneath surface of 
water. 

g. Belly strongly pigmented or somewhat pigmented, in spirits looking 
white and viscera not visible or but slightly visible; tail covered with 
large prominent dark spots; transformation 2.5-35 mm - 
h. Nostril equidistant eye and snout, av. 1.15 ; spiracle equidistant vent 
and snout, nearer eye (1.5) than vent, intestines show through belly 
but not quite like R. piptens; median space between second upper 
row of teeth 1.15-1.5 times in length of cither lateral part; third 
lower row 0.40-0.66 of the second lower row; spotting not so promi- 
nent as in R. capto but heavier than in R. pipens group; inadequate 
characterization. Southwestern Ohio to Kansas and Oklahoma and 
northeastern Louisiana. Rana arcolata circulosa 

hh. Nostril 1.0-1.5 fl earer eye than snout, av. 1.15; eye 1.0-1.3 nearer 
spiracle than snout, av. 1.11, median space between the second up- 
per row of labial row 1.0-1.0 times the length of either lateral part 
of this row, third lower row 0.33-0.66 shorter than the first or 
second rows, tail covered with large prominent dark spots; belly 
strongly pigmented (in spirits it looks white), viscera not visible; 
depth of tail in length of tail i.fr 3.5, av. 3.0. North Carolina to 
Florida. Rana capita 

Plates XI-i,i; XV~ } -8 

gg. Belly not strongly pigmented, in spirits dark, viscera shows through 
skin, tail in R. piptens group with fine flecks or small spots, or in R. 
palustris heavily washed with purplish or purplish black, transforma- 
tion sizes averaging smaller, 18-31 or 31 mm. [Rana aurora Jraytonti. 
Body 1. 1-1.1 in length of tail; depth of tail in length of tail 1.5-3.1 in 
its length. (See T. I. Storer's description.)] 

h. Median space in second upper labial row 1-4 times the length of 
either lateral part; third row of lower labial teeth 0.33-0.66 shorter 
than the first or second rows, usually at least 0.50, eye nearer the 
snout than spiracle or equidistant, nostril nearer eye than tip of 
snout; depth of tail in length of tail 1.3-3.1, av. 1.7; spiracle 1.5-1.8 
nearer eye than snout, av. 1.63. Hudson Bay to Texas and eastern 
states. Rana palustris 

Plate XI-I4 



64 HANDBOOK OF FROGS AND TOADS 

hh. Median space 0.5-1.0 or 1.0-1.5 times either lateral part of the 
second upper row, third row of labial teeth o 2.2. or 0.185-0.33 
shorter than the first lower row, depth of tail in length of tail 
1.0-1.8 or 1.7-3.4. 

i. Median space of second upper row 0.5-1.0 times either lateral 
part, third lower row of teeth 0.185-0 33 shorter than the first 
lower row, nostril to snout equal to nostril to eye, eye 1.15-1.3 
nearer tip of snout than spiracle, av. 1.1; body length in tail 
length 1.35-1.66, av. 1.5, mouth 0.9-1.6 larger than internasal 
space, depth of tail in length of tail 1.0-1.88, av. 1.65, tail crest 
usually with wide prominent dark spots, greatest length of tad- 
pole 74 mm ; spiracle 1.1-1.86 nearer eye than snout, av. 1.46. 
Southeastern states to Texas, Oklahoma, and Indiana. 

Rana piptms sphcnoccphala 
Plate XI-9,io 

ii Median space 1.0-1.5 times either lateral part, third lower row 
o 11 shorter than the first lower row, nostril i.i-i 5 nearer the 
eye than snout, eye nearer spiracle than snout 1.1-1.3; body 
length in tail length 1.3-1.0, av. 1.7, mouth larger than inter- 
nasal space, depth of tail in length of tail 1.7-3 4> av - 3-> ta il 
crest usually translucent with fine spots or pencilmgs, greatest 
length of tadpole 84 mm., spiracle i 4-1.86 nearer eye than 
snout, av. 1.59. North America east of Sierra Nevada and south- 
ward into Mexico. Rana p. ptptent 

Plate XI-n 

ff. Tadpoles 83-141 mm , tadpoles usually winter over at least one season; 

transformation sizes 18 59 mm. (except R. vtrganpes 15-35 mm > possibly 

R. Jishcri), tadpoles usually with strongly pigmented bellies, viscera not 

plainly showing through the skin (in spirit specimens). 

g. Tadpole with prominent continuous black crest margins and a black 

musculature band, belly bluish; tadpoles to 95 mm , young tadpoles 

black with transverse yellowish band on the body, spiracle 0.86-1.1 

nearer vent or base of hind legs than snout, average i.o, spiracle 0.85- 

1.1 nearer eye than base of hind legs or vent, i.e., usually equidistant, 

eye equidistant from spiracle and tip of snout, muciferous crypts very 

distinct, spiracle below lateral axis; tail tip acuminate, second upper 

labial row in upper fringe 1/3 to 1/4, upper fringe distinctly greater 

than horny beak, median space between two parts of second upper 

labial row i to i\% of either lateral part, third lower labial row equal 

to horny beak, third lower labial row longer than single row of lower 

papillae, third lower labial row 1/4 to 1/5 shorter than first lower row. 

Eggs not described. South Carolina to Florida and Louisiana. 

Rana beckscheri 
Plates XI-7,8; XIV-i,i; XCVIII-i,6 



GENERAL ACCOUNT 65 

gg. Tadpole without black crest margins or lateral band, belly white, car- 
tridge buff or yellow to maize yellow, no transverse yellow band in 
young tadpoles, spiracle nearer vent than snout 1.1-1.8; spiracle to eye 
rarely less than 1.2.5 greater than spiracle to vent, eye nearer tip of snout 
than spiracle 1.0-1.4, second upper labial row to upper fringe 1/4 to 1/15, 
upper fringe equal to or slightly greater than horny beak, median space 
in either lateral part ij^ to u; third lower labial row much less than 
horny beak, third lower row much shorter than or equal to single row 
of lower labial papillae. 

h. Tadpoles to 140 mm., eye well above lateral axis, muciferous crypts 
indistinct; spiracle just below lateral axis, spiracle 1.08-1.44 nearer 
base of hind legs or vent than tip of snout, av. 1.2.6; depth of tail in 
length of tail 2-4-3. 5 av * x -8> ta ^ C1 P obtuse; second upper row in 
upper fringe 1/4 to 1/7, median space in second upper row \Yi in 
either lateral part, third lower row in first lower 1/4 to 1/5 shorter; 
teeth 1/3, rarely 3/3. Transformation size 43-59 mm. Egg mass, 
surface film. North America east of Rockies. Rana catesbciana 

Plate XCIV-3 

hh. Tadpoles to 83-110 mm.; eye on or just above lateral axis, tail tip 
acute or acuminate (rounded in R. fishcrt)\ teeth, 1/3, second upper 
row fringe 1/6 to 1/15, median space in second upper row 1.5-11 in 
either lateral part, third lower row in first lower row 1/2. to 1/3 
shorter. 

i. Depth of tail in length of tail 1.45-1.8, average 1.7; tail tip acumi- 
nate, dorsal crest equal to or less than tail musculature, muciferous 
crypts indistinct, spiracle 1.08-1.44 nearer vent than snout; mouth 
in interorbital distance 1.5-2.. 37, av. 1.94, internasal space in in- 
terorbital distance 1.8-1.6, av. 1.16; second upper row 1/6 to 1/8 
of the upper row, median space of the second upper row 2.% to 
4!^ times either lateral row, third lower row i 5 less than horny 
beak, much shorter than single row of lower labial papillae and 
1/2. shorter than first lower row of teeth, first row of lower teeth 
equal to horny beak. Transformation size 31 or 37 to 48 mm.; egg 
mass, a surface film. Georgia to Florida to Louisiana. Rana grylto 

Plates XI-5,6, XIV -5 

ii. Depth of tail in length of tail 1.5-4.7, avs. 3.1-3.87, tail tip acute, 
dorsal crest less than tail musculature, muciferous crypts distinct, 
species 1.07-1.8 nearer vent than snout, mouth in interorbital dis- 
tance 1.3-1.8, av. 1.5, internasal space in interorbital space i 15- 
i.o, avs. 1.6-1.75; second upper row 1/6 to 1/15 of the first upper 
row; first lower row of teeth equal to or greater than horny beak, 
spiracle 1.1-1.8 nearer vent than snout. 



66 HANDBOOK OF FROGS AND TOADS 

j. Tadpoles to 99 mm., transformation size to 19-38 mm.; depth of tail in 
length of tail 3.1^4.7, av. 3.87; spiracle just touches lateral axis; eye just 
above lateral axis, spiracle 1.06-1.38 nearer eye than base of hind legs 
or vent, av. 1.2.4, spiracle 1.15-1.6 nearer eye than vent, 1.45, mouth in 
interorbital distance 1.3-1.75, av. 1.55, width of body in its own length 
1.3-1.1, av. 1.55, third lower labial row of teeth 1.15 less than horny 
beak, about equal to single row of lower labial papillae, 1/1 shorter than 
first lower row, sometimes a row of inner papillae below the third lower 
row of teeth, median space in second upper labial row 3.5-4.5 times either 
lateral portion; second upper row 1/6 to 1/15 of the upper fringe, belly 
straw yellow, colonial buff, or deep colonial buff, tail with round car- 
tridge buff or pinkish cinnamon spots, no black line in dorsal crest as in 
R. grylio or R. virgattpcs. Eggs in a compact submerged mass. Hudson 
Bay to Minnesota, New York to New England. Rana scptcntrionalts 

Plates XI-n; XIV- 3 

j). Tadpoles to 91 mm , transformation sizes at 15-38 mm., depth of tail in 
length of tail 1.5-3 7> spiracle just below lateral axis, eye on lateral axis, 
third lower labial row of teeth 1.5- 1.2.5 ^ ess than horny beak, much 
shorter than single row of lower labial papillae, almost 1/2. shorter than 
first lower row, median space of second upper row 6-n times the length 
of either lateral portion; second upper row 1/15 to 1/15 of the first upper 
row. 

k. Spiracle nearer vent than snout 1.35-1 8, mouth 1.3-1.8 times in inter- 
orb'tal distance, av. i 5, width of body in its own length 1.15-1.7, 
av. 1.47; belly deep cream color, tail green, mottled with brown and 
covered with line yellow spots. Egg mass, surface film. Canada to 
Louisiana to Florida to New England. Rana damttans 

Plates XI-3,4, XIV- 7 

kk. Spiracle nearer vent than snout 1.07-1.45, av. 1.13-1.3, mouth i.o- 

1.37 in interorbital distance, av. 1.11. 

1. Teeth 1/3 or 1/3 , no inner papillae or few inner papillae from end of 
the upper row to the end of the lower labial row, no row of finer 
papillae below third lower row of teeth, spiracle nearer eye than 
vent 1.1-1.75, ' lv - * 43' nostril nearer eye than snout 1.1-1.9, av - 
1.5, tadpoles medium ^our material, 41 mm., Stanford University 
material, 83 mm.), width of body in body length 1.6-1.0, av. 1.77; 
second upper row 1/6 to i /8 of the first upper row, belly pure white 
or pale cinnamon pink, tail musculature with black clusters out- 
lining cartridge buff areas, upper tail crest sometimes reticulated 
with black dots. Lower crest except for caudal half free of spots. 
Eggs unknown. Transformation size unknown. Nevada. 

Rana fishcri 
Plate XIII-i 



GENERAL ACCOUNT 67 

11. Teeth 1/3 or 1/3 ; four to six rows of inner papillae 
from end of first upper row to end of lower labial 
row; a row of heavy inner papillae below the 
third lower row of teeth; spiracle nearer eye than 
vent i. 41-1,82., av. 1.62.; nostril nearer eye than 
snout, 1.0-1.42., av. 1.2.; width of body in body 
length 1.45-1.86, av. 1.6, tadpole large (92. mm.); 
second upper row 2.715 or 1/15 of the first upper 
row or second upper row absent, belly pale chal- 
cedony yellow, sulphur yellow, vinaceous, pale 
grayish vinaceous or vinaceous-buft"; upper fail 
crest with a black line or row of large black spots, more 
prominent than in R. grylio, middle of musculature 
ivtth another black line, tail dark ivitb pale chalcedony 
yellow spots. Transformation size 2.5-35 mm - Eggs 
a submerged mass. New Jersey to Okefinokcc 
Swamp, Ga. Rana vtrgattpes 

Plates XI -13, XIII i 

dd Papillary border on side of labium without an emargination, tadpoles 
13-50 mm. length; labial teeth 1/3 or z/2.. HYLIDAE 

e. Labial teeth z/i, tip of tail normally black (sometimes lost), median 
space of second upper tooth row wide (like Rana) and 1.33 greater 
than cither lateral portion, horny beak 1.1-1 35 m first upper row, 
eye dorsal just inside the lateral outline in dorsal aspect (more like 
Ranids). Eye i.o 1.66 (av. i.zz) nearer tip of snout than spiracle; 
depth of tail in length of tail 3.2.5-5 o, av. 40, suborbital region 
oblique, not vertical; spiracle to eye usually equals distance from 
spiracle to vent; spiracle plainly showing from dorsum, sptracular tube 
tn Itfe stands out at an angle from the body and opening tf apart from the body 
proper; papillary border does not end above the end of the upper row 
(like Rana), eggs single, occasionally a mass. Acrts 

Plates X-n; XII 6, XLV 4, XLVI- 9 

ee. Labial teeth usually 1/3, rarely 1/1, tip of tail not normally black, 
median space (sometimes absent) of second upper tooth row narrow, 
1.6-4.0, rarely 4 5-7.0 in either lateral portion, j e., smaller than 
cither lateral part, horny beak or mandible 1.3* 2..3 in first upper row, 
eye lateral visible from ventral as well as the dorsal aspect; eye i.o- 
i 75 nearer spiracle than tip of snout; depth of tail in length of tail 
1.6-4 4> av - 2 - I ~"3 '55 i suborbital region vertical; spiracular tube in life 
parallel with opening at inner edge closely connected with or near to 
body proper. HYLIDAE exclusive of Acris 



68 HANDBOOK OF FROGS AND TOADS 

f. Labial teeth occasionally, or frequently 1/1; tadpoles 13-33 mm - 

g. Tail musculature with black-brown lateral band with light area below; tail 
crests clear with fine elongate fleckings, single row of papillae on lower 
labial border below second lower row of teeth; upper fringe somewhat 
angulate in middle (like H. femoralis); spiracle 1.4-1.9 nearer base of 
hind legs or vent than tip of snout, av. 1.41, nostril to eye 1.37-1.0 in 
nostril to snout, av. 1.6; median space between the second upper row 
of teeth 1^-5 in either lateral portion; ends of second lateral row ex- 
tending not at all or slightly beyond the end of the upper fringe; horny 
beak about 1.5-1.6 in upper fringe. Eggs an irregular mass. 

Pseudacns nigrita triseriata 

gg. No black-brown dorsal and white or light lower band on tail musculature; tail 
fins clear except where heavily pigmented with purplish black blotches 
on the outer edge of the tail fin; two rows of papillae on lower labial 
border below second lower row of teeth, upper fringe not perceptibly 
angulate in the middle; spiracle 1.0-1.6 nearer base of hind legs or vent 
than tip of snout, av 2..i6, nostril to eye i.o in nostril to snout; 
median space between second upper row of teeth 1-3 in either lateral 
row; ends of second lateral row extending beyond upper fringe for 1/4 
to 1/6 of either lateral portion, horny beak 1.75-1.15 in length from 
one end of second upper lateral row to end of other lateral row. Eggs 
single. Hyla cructfer 

Plates X-3,XII-i 

ff . Labial teeth 1/3 . 

g. Third row of labial teeth short, shorter than horny beak or 0,10-0 40 
of the first lower row in length, first upper row very angulate (i form), 
somewhat angulate (i forms), not angulate (3 forms), no flagellum 
ordinarily present yet some have pointed tails, tadpoles 13-50 mm., 
light papillary development, lower labial corner not with three or 
four strong rows of papillae, one or two rows of papillae below third 
lower row of teeth or none (Ji . crucifer)', the papillae extend above and 
beyond the end of the upper fringe for about 0.14-0.185 of the length 
of the upper fringe. 

h. Tadpoles 13-33 mm., eye equidistant between spiracle and tip of 
snout (o.8--i.3 times); spiracle 1.1-1.6 nearer vent or base of hind 
legs than tip of snout, spiracle i .0-3 .1 nearer eye than vent, papillae 
extend above the fringe for 0.16-0.15 of the length of the first 
upper row. 

i. Musculature with no distinct brown lateral band with light area below; 
crests usually heavily pigmented with purplish black blotches 
on outer rim, nostril to eye in nostril to snout 2. o, spiracle to 
vent 1.0-1.6 in spiracle to tip of snout; no papillae below third 



GENERAL ACCOUNT 69 

lower labial row of teeth thus appearing as a goatee; median space between 
second upper labial row 1-3 in either lateral portion; horny beak in first 
upper row 1.75-1.15. Eggs single, submerged. End of second upper row 
extends beyond end upper fringe 1/4 to 1/6 width of second lateral row; 
upper fringe not angulate in middle; median space in second upper row 
1.4-1.5 in horny beak; third lower row in first or second lower row 3-5 
times; second lower row distinctly longer. Hyla cructfer 

Plates X- 3 , XII -i 

ii. Musculature with a distinct brown lateral band above with light area below > i.e., 
bicolored; crests usually clear with fine scattered fleckings, sometimes with 
fleckings gathered nearer outer rim, one or two rows of papillae below 
third lower labial row of teeth (rarely absent), spiracle to vent in spiracle 
to tip of snout once or i.i -i.o or 1.1 times, one or two rows of papillae 
below the third lower labial row of teeth; median space between second 
upper row 1.6-5.0 times in either lateral portion, horny beak in upper 
fringe 1.3-1 3, end of second upper lateral row to end of first, slightly or 
distinctly shorter, or rarely slightly beyond, first upper row very angulate 
(i form) somewhat angulate (3 forms), not angulate (2. forms) in middle; 
median space (absent in i form) of second upper row 1.5-3.5 * n horny 
beak, third lower row of teeth from i 1-4 times in first or second lower 
row; second and first equal (except one with first considerably shorter 
and one equal or first 1.1-1.1 shorter). Eggs in a mass except P. oculans 
where single. Pseudacris 

j. Tadpoles 15^-13 mm., adults 11.5-17.5 mm , eggs single; dorsal crest 
to vertical midway between spiracle and eye, spiracle equidistant from 
eye and vent; median space 3.5 in horny beak, horny beak i.o m upper 
fringe, horny beak i.o in first or second lower rows of teeth, third 
lower row of teeth 3-4 in first or second lower rows; third lower row 
4.0 in first upper row, first upper row somewhat angulate in middle; 
mouth and internasal space equal, spiracle 1.7-1 i nearer vent than 
snout, one row of papillae below the third lower row of teeth; dorsum 
of body tn Itfe with definite scattered black spots; musculature with three 
bands, apricot buff (light), chestnut brown (dark), martius yellow 
(light). Pseudacns oculans 

Plates X-8; XII-8 

jj. Tadpoles 2.3-36 mm.; adults 19^-48 mm., eggs in a mass, dorsal crest to 
vertical of spiracle or within i or z mm. of such vertical; spiracle to 
eye 1.0-3.0 or 3.1 nearer eye than spiracle to vent (one form sometimes 
equidistant); median space usually present, 1.5-3.0 in horny beak; 
horny beak 1.3-1.3 in first upper row; horny beak in first or second 



7 o HANDBOOK OF FROGS AND TOADS 

row 1111, rarely over i.o times, third lower row of teeth in first or second 
lower row i i- 1.8, rarely 3.0 3.1 times, third lower row in first upper row 
1.6 -3.8 occasionally 4 o. Pstudacns (exclusive of P. oculans) 

(k. Adults 15 -48 mm , first and second lower tooth rows equal and united 
at their outer ends, median space of first upper tooth row very angulate; 
musculature depth 3.0 in tail depth, median space of second upper row 
i. 6-i 7 in either lateral part, median space i 5 in horny beak, horny 
beak 1.1 2. 3 in first upper row, third row 1.3 in horny beak, mtcrnasal 
space i. 6 -i. 7 in mterorbital space, tail depth greater, brown gray 
dorsally, with median mterorbital rectangular darker area and a crescent 
at rear of each nostril, belly white. (After Bragg )) Pscudacns stricken 

kk. Adults to 19 36 mm , musculature depth in tail depth i 1-1 9, median 

space of upper labial tooth row somewhat angulate or not angulate, 

median space of second upper row 1-5 in cither lateral portion (absent 

in i form, sometimes absent in another), median space of second upper 

row in horny beak 3 o or more, horny beak i 3-2. o in first upper row; 

third row 13 3 o in horny beak, first and second lower rows equal, or 

first, ri i i, or considerably shorter, free at ends (in one form almost 

united), internasal space in interorbital 1.33 i 4, usually i.o or more, 

rail depth in tail length usually 11-40, rarely 4 4. 

(1 Tail depth in tail length 1115, musculature of tail 17 3.0 in tail 

depth, eye to spiracle 1.5 in eye to tip of snout, internasal space i 8 

in mterorbital space, internasal space i.o in mouth, nostril equidistant 

from eye and snout or i 15 nearer eye than tip of snout, iirst upper 

row not angulate in middle, median space of second upper row i o- 

i 3 in cither lateral part or 3.0 in horny beak, horny beak i 6-1 8 in 

first upper row, or i 6 in first or second lower row, third lower row 

1.5 i 6 in horny beak or 16 1.8 in first or second row, first and second 

lower rows equal and almost united on their ends, black-brown above, 

under parts bronzy black, black along side, body transparent. (After 

Green )) Pseudacns bracbypbona 

11. Tail depth 1.4 5.0, seldom under i 5 in tail length, musculature 1.1- 

i 6 in tail depth, eye to spiracle 0.8 1.36 in eye to snout, internasal 

space 1.3 1.4 in mterorbital space, internasal space about equal or i.i 

or i. i~i 7 in mouth, nostril to eye 1.1-1 o in nostril to snout. 

m. Upper fringe of teeth not angulate, no median space in second 

upper row or median space 1.8-1.0 in either lateral part or i.o in 

horny beak, horny beak in upper first row of teeth or fringe 1.3- 

1.7, first and second rows unequal, first considerably (1.1-1.5) 

shorter than second row; horny beak 1.1-1.5 in second row and 



GENERAL ACCOUNT 71 

1.1-1.4 greater than first row, horny beak 1.3-1 o in upper fringe; 
third lower row 1.5-3.0 in horny beak; one row of papillae under 
third lower row; spiracle nearer eye than vent, tail depth in tail 
length 1.4, musculature of tail 1.6 in tail depth, third lower row 
1.0-1.3 in first lower row, 1.3-3 1 in second lower row, gray olive 
or grayish brown tadpoles. Pswdacrts n. clarkti 

Plate XIII 4, L- 5 

mm. Upper fringe of teeth somewhat angulate, median space of second 

upper row 1-7 in cither lateral portion of this row (occasionally 

absent in P. //. ftnawtri), or 3 or more in horny beak, first and 

second lower tooth rows equal or first 1.1-1.1 shorter than second 

row, third lower row 1.3 1.8 m horny beak, horny beak 1.5 i.o 

in upper fringe or 1.3-1.1 in first or second lower rows; spiracle 

to eye 1.0-1.7 nearer than spiracle to vent; tail depth 15 3.15 in 

tail length, musculature of tail 1.1 1.5 in depth of tail, third 

lower row i 0-3.0 m first and second lower row. 

n. Two rows of papillae under third row of teeth, median spate 

of second upper row sometimes absent, if present ij *j or 3-7 

m lateral part, horny beak i o-i i in first or second lower 

rows of teeth, third lower row i 3-1.4 in horny beak or 3 4 

m upper fringe. Puudacns n fenarum 

Plate XII 5 

nn. One row of papillae under third lower row of teeth; median 
space of second upper row i 5 in either lateral row, horny beak 
i 3- j 5 in first or second lower rows of teeth, third lower row 
i 8 in horny beak or 3 o in upper fringe. 

Ptcudacris n trtscnata 

[The key to PtcuJacns may be worthless, and the characters 
given here may be extremely variant.] 

hh. Tadpoles 35-50 mm., tadpoles not bicolorcd (brown above and white 
below) but green, citrine tadpoles or scarlet- or orangc-uilcd tadpoles; 
high-crested tadpoles, eye 1.0-1.75 nearer spiracle than snout, spiracle 
i o-i 6 nearer vent than tip of the snout, spiracle i 15-1 5 nearer eye 
than vent or base of hind legs, papillae extend above the upper fringe 
for o 14-0 185 of the length of the upper fringe. 

i. Tadpoles 50 mm. in length, body in tail 1.3-3 1 5 av - M depth of 
body in width of body o 83-1.0, av. o 9, depth of tail 10 14 mm , 
beautiful green tadpoles, young tadpoles with a black saddle spot on 
the back of the musculature near its base and with a light line from 
eye to tail, one row of papillae below lower third labial row, papillae 
extending above first upper row for 0.15-0.185 of the fringe's length, 



72 HANDBOOK OF FROGS AND TOADS 

dorsal crest extending to a vertical halfway between eye and spiracle. 
Eggs single, submerged. North Carolina to Louisiana. Hyla gratiosa 

Plates X-5,6, XII-i, LXIX- 5 



ii. Tadpoles 35-45 mm., body in tail 1.1-1.0, av. 1.6, depth of body in 
width of body 1.0-1.8; depth of tail 5-9 mm., no black saddle spot in 
young tadpoles. 

j. Tadpole small (35 mm.), dorsal crest extending to vertical halfway 
between spiracle and the base of the hind legs; depth of tail in tail 
length 2.. 5-3. 5, av. 30, nostril to eye 1.1-1 1 in nostril to snout; 
mouth in interorbital space 1.33-1.6, mternasal space in interorbital 
distance i 33-1.1, eye just touches lateral axis or is below it, beak in 
first upper row 1.5-1.7, papillae extending beyond the end of the upper 
row 0.15-0.185 of the length of the upper row, two rows of papillae 
below third lower labial row, median space between second upper 
labial row 1.15-10 in either lateral portion; third lower labial row 
o.io 0.11 of the first lower row, first row of lower labial teeth 1.0-1.5 
times the horny beak. Eggs strewn in water among sphagnum (Noble 
and Noble). New Jersey to South Carolina. Hyla andtrsomt 

Plate X-i,XII-3 

jj Tadpole medium (40 and 45 mm ), dorsal crest extends ahead of 
spiracle or to eye, depth of tail in tail length i 5-3.1, av. 1.75, nostril 
to eye 1.0-1.7 in nostril to snout, mouth in interorbital space i 4-1.0, 
internasal in interorbital space 1.15-1.0, eye on lateral axis, papillae 
extending beyond end of first upper row 1.4-1.5 of the length of the 
first upper row, median space in second upper labial row 3-5 in either 
lateral portion, third labial row 0.15-0.40 of the first labial lower row; 
first row of lower labial teeth i 0-1.3 greater than the horny beak, 
k. Dorsal crest to the vertical halfway between spiracle and the eye, 
depth of body in body length i 7-1.5, musculature of tail in depth 
of tail 1.75-1.4, av. 1.9, spiracle 1.4-1.3 nearer eye than vent; 
mouth 1.0-1.4 larger than internasal space, av. 1.15, two rows of 
papillae below the third lower row of labial teeth, papillae extend 
beyond the end of the first upper row 0.11-0.15 of the length of this 
fringe, horny beak in upper row 1.0^-1.3; third labial lower row 
0.15-0.40 the length of the first lower row. Eggs surface or sub- 
merged irregular mass. Eastern Maryland to Florida to Texas to 
Illinois. Hyla cmcrca 

Plates X-i; XII~4 



GENERAL ACCOUNT 



73 



kk. Dorsal crest extending to the vertical of the posterior edge 
of the eye; depth of body in body width 1.5-1.0; muscula- 
ture in depth of tail 1.3-1.8, av. 1.5, spiracle 1.6-1.75 nearer 
eye than snout; mouth 1.0-1.1 larger than internasal space; 
one row of papillae below the third lower labial row of 
teeth, papillae extends beyond the end of the first upper row 
0.14-0.10 of the length of this upper fringe; horny beak in 
upper fringe 1.4-1.8; third labial lower row 0.15-0.33 the 
length of the first lower row. Eggs loose irregular mass. 
Vancouver to Lower California and east to Utah and Arizona. 

Hyla regtlla 

gg. Third row of labial teeth long, longer than horny beak, or 0.75-1.00 of 
the first lower row in length; first upper row very angulate in middle; 
flagellum on tail; tadpoles 31-50 mm,, heavy papillary development, 
lower labial corner with three or four rows of papillae; two more or less 
complete rows of papillae below third row of teeth (except in Hyla 
aremcolor), papillae extend above and beyond the end of the upper fringe 
for about 0.3-0.4 of its length. 

h. Third lower labial row 0.8-1.0 of the length of the first lower row; 
dorsal crest extends to the vertical halfway from hind legs to spiracle, 
to spiracle, or halfway from spiracle to eye, dorsal crest equal to, 
greater, or less than depth of tail musculature; tadpoles 36-50 mm.; 
red may be present in the tail, tail crest distinctly or more or less 
clear of spots next to the musculature, tail heavily blotched with 
dark blotches or spots. 

i. Median space between lateral upper rows 5.0-10.0 times in either 
lateral row, spiracle 1.44-1.5 nearer eye than vent, width of body 
in its own length 1.6-1.1, eye 1.0-1.7 nearer spiracle than tip of 
snout; tail sometimes suffused with coral red or coral pink, 
j. Median space between lateral upper rows of teeth contained 
6.a-io.o in either lateral row, first and second lower rows of 
teeth 1.4-1.6 greater than horny beak; mouth equal to inter- 
nasal space; depth of tail in length of tail 1.6-1.75, av. 1.15; 
muscular part of tail in depth of tail 1.8-1.3, av. 1.1; depth 
of body 1.33-1.1 in body length, av. 1.68; dorsal crest usually 
equal to or greater than musculature depth; center of belly 
solid sulphur yellow; tail )-j-banded; light lateral band bounded 
above and below by a brown band, flagellum clear of pigment; 
body olivaceous black. Eggs a surface film. Virginia to Florida 
to Texas . Hyla fcmorahs 

Plates X-4; XII-i 3 ; LXVIII- 3 



74 HANDBOOK OF FROGS AND TOADS 

jj. Median space between lateral upper rows contained 5.0-10.0 times 
in either lateral row, first and second lower rows of teeth 1.5-1.0 
greater than horny beak; third lower row of teeth may be equal to 
or slightly shorter than the first lower row; no or few papillae be- 
neath the third lower row of teeth, surely not a complete row; 
mouth in internasal space 0.83-1.7, av. 1.3; depth of tail in length 
of tail, x. 85-5. 15, av. 3 .75 ; musculature of tail in depth of tail 1.18- 
1.76, av. 1.6; depth of body in body length 1.75-1.3, av. i.o; dorsal 
crest halfway to, or to, vertical of the spiracle; dorsal crest less than 
the musculature; flagellum or tail tip spotted; body greenish olive 
or deep olive; center of belly solid pale cinnamon pink. Eggs single, 
submerged (Atsatt and Storer). Western Texas (Devil's River, Fort 
Davis Mountains, etc.), Utah, California, Mexico. Hyla arenicolor 

Plate LX-6,7 

ii. Median space between lateral upper rows contained 3.15-5.0 times in 
either lateral row; spiracle 1.11-1.5 nearer eye than vent; eye about 
equidistant between spiracle and tip of snout; internasal space in 
mouth 0.7-1.0, dorsal crest extends to vertical of spiracle or halfway 
between eye and spiracle; dorsal crest equal to or greater than muscu- 
lature depth; muscular part of tail in depth of tail 1.71-1 9, av. 1.8; 
depth of tail in length of tail 3.1-4.0; width of body in its own length 
i .3-1 .7; no lateral bands in tail; tail more or less scarlet or orange vermilion 
with black blotches more prominent near the margins of the crests; bodies 
olive-green; belly conspicuously white or very light cream. Eggs a 
surface film. Minnesota to Texas to Maine to Florida. 

Hyla v. versicolor 
Plates X~9,io; XII-n 

hh. Third lower labial row 0.75 of the length of the first row, dorsal crest 
extends to the vertical of the posterior edge of the eye; dorsal crest usu- 
ally less than depth of the musculature; tadpoles to 31 mm.; width of 
body in its own length 1.7-1.1, av. 1.875; depth of tail in length of tail 
1.1-3.3, av. 1.8, third lower row of teeth not equal to first lower row; 
median space between lateral upper rows contained 3.15-5.0 times in 
length of either lateral row; papillae extend above and beyond the ends 
of the upper fringe for 0.3-0.33 of the upper fringe, horny beak in upper 
fringe 1.8-1.0; no bands or red in tail; tail crest clear, uniformly sprinkled 
with distinct black dots; body greenish (like H. cinerea or H. gratiosa); 
belly testaceous or chalcedony yellow. Eggs single, submerged. Texas 
to Indiana to Florida to Virginia. Hyla squirella 

Plates X- 7 ; XII- 9 



GENERAL ACCOUNT 



75 



Development and transformation: 

Some frogs have limited breeding pc 
riods and other species may breed al- 
most any month in the year. The 
males usually precede the females to 
the water and croak vigorously dur- 
ing breeding time. The male with ils 
forearms seizes the female. In almost 
all frogs the eggs are fertilized just at 
or slightly after the extrusion of the 
eggs. At first no envelopes about the 
eggs are apparent, and the egg mass 
may feel soft and sticky. After a few 
minutes this substance absorbs water, 
and each egg is then revealed with its 
vitelline membrane and one or more 
jelly envelopes. 

The eggs hatch in 3 to 25 days, 
depending on temperature and other 
conditions. At hatching the larva has 
a distinct neck, with a prominent 
head and body. The tail is very small 
or absent. On the ventral side of the 
head is an invagination or depression 
which is to be the mouth. Behind this 
comes the ventral adhesive disk or 
disks, which help the little creature 
to attach itself to the egg mass or to 
hang itself upon some plant. In front 
of the mouth are two deep, dark pits 
which later become the nostrils. On 
either side of the head appear swell- 
ings which become the external gills, 
The eyes do not yet appear. 

As development goes on, the exter- 
nal gills appear as branched organs : 
two or three on a side; the eye shows 
as a ring beneath the skin; and the 
tail grows and presents a middle mus- 
cular portion where the muscle seg- 
ments clearly show. This middle part 
supports a thin, waferlike tail fin the 
parts of which are called the lower 




? I 



Plate XV. Development of the gopher 
frog, Rana capita, i. Egg mass (X%)- 2. 
Eggs (X^)- 3-7- Tadpoles (X%)* U- 
Lateral lines and spiracle; 5, with two legs; 
6, with three legs; 7, with four legs.] 8. 
Transformed frog (X%) 9- Adult 



76 HANDBOOK OF FROGS AND TOADS 

and upper crests. The nasal pit shifts in position and becomes the nostril, and 
the vent opens. The mouth appears, and dependence on the yolk of the belly 
ceases. Soon the external gills begin to disappear, a lateral dap or fold of skin 
connects the head with the body, and the neck region disappears. Beneath 
this fold internal gills develop. Usually on the left side but on the middle line 
in the belly in ribbed frogs and narrow-mouthed toads, the flap does not close 
completely, but leaves an opening, the spiracle. The water passes into the 
mouth over the internal gills and out of this hole. On the mouth a membra- 
nous, fringed lip, with upper and lower portions (labia) comes into being. 
At the portal are horny jaws or mandibles. On the upper and lower portions 
are ridges of horny teeth. The eyes are no longer covered, pigmented rings but 
are now at the surface. The intestine has become much elongated and coiled 
and in some can be seen through the skin. The skin of the back and head 
comes to have a series of sense organs (or lateral line dots). 

The buds of the hind limbs begin to appear. The fore limbs start to develop 
beneath the skin. When the hind limbs have reached considerable size, the 
left arm comes out through the spiracle, or the skin breaks down and the right 
arm appears through the skin or the skin weakens for its egress. It is held that 
the kft arm normally comes out first, but often the right arm appears first. 

The process of transformation continues. The tail crests decrease in size and 
the creature begins to live on its tail that is, to absorb it. The gills vanish, and 
the lungs begin to serve as the sole respiratory organs, if the skin be not con- 
sidered. The tadpole appears more and more at the surface or near the shore. 
The eye assumes eyelids. The tadpole mouth fringe, with its horny jaws and 
horny teeth, is discarded, and a true frog mouth begins to appear. The long 
intestine becomes wonderfully shortened, for a carnivorous diet, and the small 
frog, with a vestige of a tail, is ready to leave the water. This process is termed 
transformation or metamorphosis. 

In the accompanying table of 51 tadpoles, wherein total length, tail length, 
body length, and range of transformation size appear, we have a surprising 
correlation in an ascending scale for all four characters. The first criterion of 
arrangement is the arbitrary choice of total length. Strictly speaking, body 
length and transformation sizes should have governed the arrangement, but 
all seem more or less correlated. The most striking point is that the body 
length of the largest tadpoles for each species falls within the range of trans- 
formation size in all except six forms. In two of these the differences are only 
2 mm. and i mm. In the spadefoot, Scaphiopus couchii, the body length of the 
tadpole is 2 mm. greater than the known size at transformation, and in Ham- 
mond's spadefoot, Scaphiopus hammondii, it is 5 mm. greater than the known 
transformation size. In Scaphiopus holbroo^ii no such condition obtains. Is 
it because these two species are desert species which breed in the most transi- 
tory places, and is there consequently a decided shrinking in body, before 
transformation, to speed development? Many have thought there is consid- 



GENERAL ACCOUNT 



77 



erable shrinkage in the body of the tadpole before transformation. This may 
be true of some species, but in the following table there is no striking decrease 
as in the paradoxical toad (Pseudis paradoxus). Rather, the uniform parallel 
in body length of tadpole and of transformed individual is surprising even to 
the present authors, who have long worked with them. 

At least 35 of the 51 tadpoles arc less than 50 mm. in total length, and of the 
26 species whose tadpoles are unknown or not minutely described at least 12 
more will doubtless fall in this group of small or medium tadpoles. In our 
descriptions we have used small, medium, and large for tadpoles. We have 
called small 25-35 mm ; medium 40-50 or 55 mm.; large 60-140 mm. (large, 
60-86; quite large, 95-100; very large, 110-145). 

In the table at least 32 species have tadpoles with body lengths of 18 mm. or 
less. Probably in time we will know that 50 of our species fall in this class, or, 
put in another way, at least two-thirds of our frog species will be found to 
transform at 18 or 20 mm. or less. Kxcept for the true frogs (Ranidae), Ham- 
mond's spadcfoot, and a toad or two (Bufo), no species grows tadpoles over 
2 inches (50 mm.) in length. 

We cannot say that the order of maximum length of tadpoles corresponds 
with the order of maximum length of adults. As would be expected, the small- 
est swamp cricket frogs and narrow-mouthed toads have small tadpoles, but 
with them appear most of the toads (Bufo) and spadefoots (Scaphiopus ex- 
cept S. hammondti). Toads and spadefoots arc much larger than the tree toads 
(Hyla and Acns), which follow with larger tadpoles. Beginning with the tree 
frogs through the true frogs (Ramdac), the order of tadpole length more or 
less corresponds to that of adult size. 

Greatest Greatest Trans- 
Greatest tail body forma- 
Name length length length lion size 



Bufo qucrdcus 
Bufo debilis 
Pseudaci is oc ulat is 
Pseudacris mgnta trisenata 
Pseudacris n. septenttionalts 
Bufo valliceps 
Bufo terrestris 
Scaphiopus couchii 
Pseudacris brachyphona 
Microhyla olivacea 
Bufo punctatus 
Microhyla carolinensis 
Scaphiopus h. hurterii 
Bufo canorus 
Bufo w. jowleri 



mm. 



22 



23 

24 
24.5 

25 

25 

26 

26.4 

27 
27 
27 



mm. 



14.2 
14.2 

13 

14.2 

ii 

15.4 
16 
16.4 
16 



mm. 



8.8 
8 

10 

9 8 



10 
IO 
10 
12 

10 



mm. 
7-8 
8-1 1 

7-9 
7.5-11 

7-5-13 

7.5-12 

6.5-10.5 

7.5-12.5 

8 

10-12 

9-1 1 
7-12 
8-10 
10.5 



7 8 



HANDBOOK OF FROGS AND TOADS 



Greatest 
Name length 


Greatest 
tail 
length 


Greatest 
body 
length 


Trans- 
forma- 
tion size 




mm. 


mm. 


mm. 


mm. 


Bufo a. americanus 


27.8 


17.6 


IO.2 


7-12 


Bufo w. woodhousii 


27 


*7 


10 


10-17 


Scaphiopus h. holbroolyi 


28 


16.4 


12 


8.5-12 


Bufo compactilis 


28 


16 


12 


12 


Bufo cognatus 


29 


i7 


12 




Pseudacrls n. clartyi 


30 


*9 


II 


8-I 3 


Hypopachus cuncus 


3 






10-12 


Bufo hemiophrys 








9-13-5 


Hyla squlrella 


3* 


20 


12 


11-13 


Pseudacrls ornata 


32 


18 


14 


I4-l6 


Leptodactylus labialis 


32 


15 


17 


16 


Pseudacrls n. feriarum 


33 


22.4 


10.6 


8-12 


Hyla crucifer 


33 


22 


II 


9-14 


Hyla fern or alts 


33 


22.5 


IO 


10-15 


Pseudacris strecl(erl 


33 


21 


12 




Hyla andersonil 


35 


23 


12 


11-15 


Bufo exsul 


35 


2O 


15 




Hyla c. cinerea 


40 


25 


15 


12-17 


Acrls g. gryttus 


42.2 


29.2 


J 3 


9-15 


Hyla wrlghtorum 


43 


26 


17 


11-17 


Bufo b. boreas 


44 






9.5-12 


Hyla v. verslcolor 


46.6 


30.8 


15.8 


13-20 


Hyla regilla 


46.6 


3 


16 


11-17 


Acris g. crepitans 


47 


32 


15 


11-15 


Rana b. boylii 


47 


29 


18 


23-30 


Hyla gratiosa 


5o 


3i 


*9 


14-20 


Hyla arenlcolor 


5 


33 


15-4 


14-16 


Rana a. aurora 


5 


3 


20 


17-21 


Rana s. sylvatlca 


50 


33 


7 


16-22 


Rana s. cantabrlgensls 


50 


3 


20 


14-22 


Ascaphus truei 


5* 


33 


18 


14-18 


Hyla baudlnii 








21 


Scaphiopus hammondii Intermontanus 


5 2 


3 


22 


20-24 


Bufo b. halophllus 


56 


33 


23 


12-15 


Bufo b. nelsoni 








I5-l8 


Rana p. sphenocephala 


62.5 


37-5 


25 


18-33 


Scaphiopus h. bombtfrons 


6 3-95 








Scaphiopus h. hammondii 


65 


37 


28 


13-23 


Rana p. pretiosa 


68.5 


43 


25.5 


16-23 


Rana cascadae 


70 




22 


13-17 


Rana b, muscosa 


70 






20-24 



Cj 

Name 


ffcNfcKAL ACCOl 

Greatest 
length 


JNT 

Greatest Greatest 
tail body 
length length 


79 

Trans- 
forma- 
tion size 




mm. 


mm. mm. 


mm. 


Rana b. sierrae 


72 


48 23.5 


21-27 


Rana palustris 


75.8 


48.8 27 


19-27 


Rana aurora draytonii 


83 


55 ^8 


27 


Rana fisheri 


83 




28-30 


Rana areolata circulosa 






30 


Rana capita 


81 


53 29 


27-35 


Rana pipiens 


84 


56 28 


18-31 


Rana damitans 


84.8 


57 27.8 


28-38 


Rana tarahumarae 


88 


56 32 


3-35 


Rana virgatipes 


92 


63 30 


23-31 


Rana septentrionalis 


99 


67 32 


29-40 


Rana hccfacheri 


95 


53-5 4i-5 


3i-49 


Rana grylio 


no 


64.4 35.6 


32-49 


Rana catesbeiana 


142 


97 45 


43-59 


Bufo alvarius 




Not described 




Bufo californicus 




Not described 




Bufo insidior 




Not described 




Hyla avivoca 




Not described 




Hyla septentrionalis 




Not described 




Pseudacris brimlcyi 




Not described 




Pseudacris n. nigrita 




Not described 




Pseudacris n. vermtosa 




Not described 




Rana a. aicolata 




Not described 




Rana scvosa 




Not described 




Rana onca 




Not described 





Journal notes: These arc customarily from our field notes. They treat of 
habitats, general habits, or breeding; sometimes of experiences ir collecting the 
frogs; of their enemies; or of their usefulness and associated ecological fea- 
tures. Occasionally, you will find comparisons with closely related forms. 
Under topics such as "Voice," excerpts from our field notes are included. In 
1925-1934 we worked on Texan forms, and in the first edition of this hand- 
book only summaries of these forms appeared. In the present edition more 
notes are added on the southwestern and western forms. These are based 
mainly on a recent (1942) 25,ooo-mile trip in that region. 

Authorities' corner: Here are given references to, or quotations from, some 
of the more important works on the various frogs. We would have liked to 
publish excerpts from all the authors cited, but space limitations forbade. Pa- 
pers that have appeared since the completion of the manuscript (1947-1948) 
have also been listed. These references are in no sense a bibliography of the 
form. 



8o 



HANDBOOK OF FROGS AND TOADS 




Figure i. American bell toad, Ascaphidae: Ascaphus truel. i. Short anal tube of female. 
2. Light band across head. 3. Rear of femur. 4. Tail of male, extending from ventral side 
of body. 5. Vent. 6. Horny excrescences (secondary breeding characters) of the male. 




Figure 2. Spadefoots, Scaphiopodidae: Scaphiopus. i. Wide interorbital space. 2. Upper 
eyelid. 3. Small round parotoids. 4. Fleshy webs. 5. Broad waist. 6. Pectoral glands. 7. 
Vertical pupil. 8. Two metatarsal tubercles, the outer large and with a cutting edge. 



Keys 

FAMILIES 



A. Male with taillikc process, female with short anal tube, no tympanum; 
pupil elliptically vertical; upper jaw toothed; short ribs present; inter- 
nasal space 4.66-8.0 in L ; light band across head; size small, i 1/8-1 
in. (2.8-51 mm.). Ribbed toads, ASCAPHIDAE 

Plate XVI 

AA. Male without tail; female without short anal tube; ribs absent; inter- 
nasal space 7.77-11.4 in L. 
B. Waist wide, body broad and thick; hind limbs short. 

C. With transverse fold of skin across head behind eyes, size small, 
3/4-1 5/8 in. (19-41 mm.); no tympanum; no parotoids; snout 
pointed, head narrow, fingers and toes without webs except 
slight in Hypopachus; eyes small and depressed. 

Narrow-mouthed toads, BREVICIPITIDAE 
Plates CXXIV--CXXVI 

CC. Without transverse fold of skin across head, behind eyes; size 
medium to large, i 1/1-8 4/5 in. (37-110 mm.), except B. debtlis 
and B. quercicus (1/3 in.), tympanum distinct or indistinct; 
parotoids present except in i species of Scaphiopus; feet with 
extensive fleshy webs; snout blunt, eyes large; head broad. 
D. Pupil vertical (by day); parotoids absent in i species, present 
but rounded and indistinct in i species, sole without sub- 
articular tubercles, skin relatively smooth; venter smooth; 
no cranial crests; males without discolored throats. 

Spadefoots, SCAPHIOPODIDAE 
Plates XVII-XXIII 

DD. Pupil not vertical, parotoids present and elevated; sole with 
subarticular tubercles; skin warty, venter usually granulated; 
cranial crests present in most species, lacking in Bufo boreas, 
B. canorus, B. compacttlis, B. dcbtlis, 5. cxsul, B. insidior, B. 
punctatus; males usually with discolored throats. 

Toads, BUFONIDAE 
Plates XXIV-XLIV 

BB. Waist narrow, body narrower and thinner; hind limbs long; no 
parotoid. 

C. Disks on digits, neither thumb nor other fingers enlarged in male. 
D. Disks transverse; venter usually smooth, subarticular tuber- 
cles saw-toothed; eggs large, male throat not discolored. 

Robber frogs, LEPTODACTYLIDAE 
(Syrrhophus, Eleuthtrodactylus) 
Plates LXXVII-LXXXIV 



82 HANDBOOK OF FROGS AND TOADS 

DD. Disks round, large or small; pupil elliptically horizontal; sub- 
articular tubercles rounded, venter usually granular or areolate; 
male throat discolored. Tree frogs, HYLIDAE 

Plates XLV-LXXVI 
CC. No disks on digits. 

D. Extensive webs on toes, thumb of male enlarged at base; 
venter smooth. Frogs, RANIDAE 

Plates LXXXV-CXXIII 

DD. Toes free; males, if different, with strong conical spine on inner 
side of first digit and another on each side of breast, and 
forehmb much enlarged. 

Robber frogs, LEPTODACTYLIDAE (Lcptodactylus, etc.) 
Plates LXXXI, LXXXII 

Spadefoots, SCAPHIOPODIDAE: Scaphiopus 

A. Parotoid absent or indistinct, tympanum indistinct, no pectoral gland. 

B. Hind limb longer (0.78-0.96 in L.); fore limb longer (1.51-1.19 in 

L.); fourth finger longer (6.18-8.0 in L ), foot with tarsus longer 

(1.57-1.76 in L), fourth toe longer (1.66-3.2.9 in L.); size small, 

i 1/1-1 1/5 in. (37-61 mm.); back uniform or with light bands. 

Hammond's spadefoot, Scaphiopus hammondii 
Plates XVHI-XX 

Limits of range and gradations need to be worked 
out. After Tanner we present this synopsis: 

(C. "Interorbital boss present, rounded, not elongate tubercle, fronto- 
parietal fontanelle present, body smooth or less rugose dorsally 
than CC; head width narrower" (width of head in L. 1.6-1.95, 
av. 1.74, eye smaller, 10.1-11.4 in L.-- A.H.W.). 

Scaphtopus h. bombtfrons 
Plate XIX 

CC. "No intcrorbital boss present. In some specimens of intermontanus 
there is a glandular interorbital elevation which resembles the 
true boss in bombtfrons" Head wider (width of head in L. 1.11- 
1.58, av. i 35; eye larger, 7.3-8.6 in L. A.H.W.). 
D. "Body rugose or with many individual prominences or warts." 
E. "No fronto-parietal fontanelle; interorbital space with 
prominent fronto-parietal bones forming ridges ... or in 
some specimens the interorbital space is filled with a glan- 
dular prominence resembling the bombtfrons species, confined 
in the main to the Great Basin area." (Hind toes longer, 
second toe 4.3-6.1 in L., third toe 3.1-4.0 in L.; fourth 
toe 1.1-1.66 in L. A.H.W.) Scaphiopus h. tntcrmontanus 

Plate XX 



KEYS 83 

EE. "A fronto-parietal fontanellc present; interorbital space 
smooth; size intermediate between bombtfrons and inttrmon- 
montanus; . . . found on Pacific Coast south into Arizona and 
Texas." (Hind toes shorter, second toe 6.1-7.3 in L., third 
toe 4.0-4.1 in L., fourth toe 1.6-3.0 in L. A.H.W.) 

Scaphtopus h. hammondn 
Plate XVIII) 

BB. Hind limb shorter (0.89-1.15 in L ), fore limb shorter (1.0-1.31 in 
L ), fourth finger shorter (8 O-IT i in L.), foot with tarsus shorter 
(1.7-135 in L); fourth toe shorter (3.01-4.0 in L.); size larger, 
i 7/8-3 1/5 m. (48-80 mm.); back greenish, more or less marbled 
with light. Couch's spadcfoot, Scaphiopus couchit 

Plate XVII 

AA. Tympanum distinct, parotoid distinct, pectoral glands present, size 
1-17/8 in. (50-71 mm.). Scapbtopus holbrookit 

Plates XXI-XXIII 

B. Head to angle of mouth smaller (3.14-3.81 in L.), width of head 
smaller (1.58-1.66 in L.), snout smaller (5.4-6.3 in L.), tympanum 
smaller (11.5-16.0 in L ). 

Hurter's spadefoot, Scapbiopus holbrookit hurtcnt 
Plate XXIII 

BB. Head to angle of mouth greater (1.93- 3.56 in L.); width of head 
greater (i 14-1 75 in L.), snout larger (5.17 -6.6 in L.); tympanum 
greater (10.1-11 o in L ); skin relatively smooth with two or more 
evident light dorsal stripes. Spadefoot, Scaphiopus h. holbrooktt 

Plate XXI 

(C. * 'Great amount of white on back, ilanks, and upper surface of 
limbs, vermiculated irregular white bands." 

Key West spadcfoot, Scaphtopus holbrookti albtts 
Plate XXII) 

Toads, BUFONIDAE. Bufo 

A. Gland on leg, fold skin on tarsus, warts at angle of mouth. 

B. Crests curved around rear of eye, size large, 3 1/5-6 3/5 in. (8a- 
165 mm.), skin smooth; head broad (1.1-1 8 in L.), color uniform, 
glands conspicuous on both tibia and femur, tympanum medium 
(11-15 m L.). Colorado River toad, Bufo alvarius 

Plate XXIV 

BB. Crests absent, gland on tibia only, pitted warts on back, tympanum 
small (14-16 in L.). 



84 HANDBOOK OF FROGS AND TOADS 

C. Dimorphic; interparotoid interval less than width of gland; parotoid 
and leg gland obscured by pattern; skin smooth; male uniform green; 
female spotted; size small, 1-3 in. (50-75 mm.)- 

Yosemite toad, Bufo canorus 
Plate XXXI 

CC. Not dimorphic; parotoids widely separated, interval greater than 
width of gland; parotoid and leg glands evident, size small, medium 
or large (41.5-115 mm.) i 4/5-5 in. Bufo boreas group 

D. Eyelid narrower (11.8-14.6 in L); eye smaller (94-11.7 in L.), 
head narrower (i 66-3.1 in L.), third toe longer (1.8-3.5 m L )i 
fourth toe longer (1.0-1.5 * n L.)- "Spread of hind foot from end 
of first toe to the fifth toe more than 36% of length" (Camp, 1917^. 

Northwestern toad, Bufo boreas boreas 

Plate XXVII 

DD. Eyelid wider (7.3-14 in L.), eye larger (7.0-10.5 in L.); head wider 

(1.4-3.0 in L.), third toe shorter (3.1-4 15 in L.), fourth toe shorter 

(i 53-3.8 in L.), spread of hind foot less than 36% of the length. 

E. Dorsum almost entirely black, size small, i 4/5-1 1/1 in. (44- 

61.5 mm ), eyelid wider (7 3-11 in L ), width of head 1.77-3.0 

in L., fourth toe longer (1.53-1.66 in L.), venter heavily spotted 

with black. Black toad, Bufo exsul 

Plate XXXV 

EE. Dorsum gray, green, or brown spotted, size medium to large, 
i 4/5-5 in. (41 5-115 mm.), eyelid narrower (9.3-14 in L.), 
fourth toe shorter (1.57-3.8 in L), venter uniform white or 
lightly or moderately spotted. 

F. Size medium to large, 1 1/5-5 * n - (60-115 mm.), hind limb 
longer (0.8-1 o in L.); head wider (1.4-1.85 in L.), fourth 
toe shorter (3.3-3 8 in L.), third toe longer (3.1-3 8 in L ). 

California toad, Bufo boreas balophtlus 
Plate XXVIII 

FF. Size small to medium, i 4/5 to 1 1/1 in. (41.5-61 mm.), 
hind limb shorter (1.0-1.1 in L.), fore limb shorter (1.86- 
1.15 in L.), head narrower (1.8-3 i m L ) fourth toe longer 
(1.57-1.94), third toe shorter (3.5-4.15 in L.), "elbows 
and knees not meeting when appressed to the sides of the 
body" (Stejneger). Amargosa toad, Bufo b. nelsoni 

Plate XXIX 



KEYS 85 

AA. No gland on leg; no fold of skin on tarsus. 

B. Femur almost entirely enclosed in body skin; vocal sac elliptical 
(sausage), i sole (metatarsal) tubercles with cutting edge. 
C Crests prominent; boss on snout; interorbital narrow (11-19 * n 
L.) less than internasal, middorsal stripe and light-bordered 
large dark spots (sometimes small spots). 

Great Plains toad, Bufo cognates 
Plate XXXII 

CC. Crests absent, interorbital broad (9.7-11.7 in L.) greater than 
internasal; drab with small, dull citrine spots 

Spadefoot toad, Bufo compactilis 
Plate XXXIII 

BB. Half or more of femur free from body skin, outer metatarsal tubercle 
without cutting edge. 

C. Parotoids oval to elongate (sometimes triangular in Bufo qucr- 
ctcus). 

D. Crests absent or obscure, parotoids broadly oval, divergent, 
interorbital broad (9.4-11 8 in L.), narrowing forward; 
snout 7.0-8.15 in L. 

Southern California toad, Bufo calif ornicus 
Plate XXX 
DD Crests present. 

E. Size small, 3/4 i 1/4 m. (19 31 mm.), vocal sac a sausage; 
mouth small (4.0 4 6 in L ), tympanum small (14- 10 in 
L.), snout long (5.7-7 o in L ), yellow stripe down mid- 
back, many red tubercles. Oak toad, Bufo qucmcus 

Plate XL 

EE. Size greater, i 5/8-4 3/4 in. (40-118 mm ), snout shorter 
(6.1-9.4 in L ). 

F. Mouth small (4.1^4 7 in L ), tympanum small (14-11 
in L.), head narrower (i 5-3.1 in L ); head shorter 
(3.4-3.9 in L.), crests, a boss, from snout to rear of 
eye, with sides parallel, snout 7.4-9 3 in L. 

Canadian toad, Bufo hcmiophrys 
Plate XXXVI 

FF. Mouth large (3.3-4.3 in L.), tympanum large (10- 
18.1 in L ), head wider (1.3 in L.). 
G. Crests prominent with knobs in rear, skin finely 
and evenly roughened with tubercles between 
larger warts, red, gray, or black; snout 6 -8 in L. 
Southern toad, Bufo tcmstris 
Plate XLI 



86 HANDBOOK OF FROGS AND TOADS 

GG. Crests low, paired spots of darker color down back, super- 
ciliary crests meeting postorbital at right angles, snout 
6.1-9.4 in L. 

H. Small uniform warts on back; several warts in each dark 
dorsal spot; dark pectoral spot, no preparotoid longitu- 
dinal crests, under parts usually unspotted; back greenish, 
size smaller, 1-3 J^ in. (51- 81 mm.); snout 7.7-9.4 in L. 
Fowler's toad, Bufo woodhousn fowleri 
Plate XLIV 

HH. Large dorsal warts; one or two warts in each dark dorsal 
spot, under parts spotted or plain. 

I. Many warts spiny, particularly on hind limbs, paro- 
toids parallel, closest together at midpoint; parotoid 
on dorsolateral line or on dorsum; no boss, preparo- 
toid longitudinal crest present, venter often spotted; 
middorsal absent or present; snout 6.1-9.1 in L. 

American toad, Bufo a. americanus 
Plate XXV 

J. Brilliant coloration; long, narrow parotoids; greater 
width between parallel cranial crests, hind limbs 
shorter, ventral granulation smooth. 

Hudson Bay toad, Bufo americanus cofei 
Plate XXVI 

II. Warts round, smaller, parotoids slightly divergent at 
rear and on lateral aspect, parotoids usually in contact 
with postorbital crest, often with boss on nostril with 
crests extending backward; venter unspotted or with 
median pectoral spot, middorsal line present, snout 
7.0-9.1 in L. 

Rocky Mountain toad, Bufo w. woodbousii 
Plate XLIII 

CC. Parotoids round or triangular; sole (metatarsal) tubercles round, small, 
noncutting; dorsal pattern without the 4-6 paired spots (of the JB. 
americanus, w. fowleri, terrestns, woodbousii, etc., group). 
D. Crest prominent, size large, -84/5 in. (50-1x0 mm.), parotoid 
subtriangular. 

E. Parotoid as large or larger than side of head, divergent, not bi- 
colored; toes 1/1-1/3 webbed, crests not trenchant and top of 
head not a deep valley, brown with some black, yellow, red, 
olive; with or without black spots on a light vertebral line; 
snout 7.3-8.6 in L. Marine toad, Bufo marmus 

Plate XXXVIII 



KEYS 87 

EE. Parotoid much smaller, not as large as side of head, not di- 
vergent, bicolored; row of light conical tubercles on side, body 
flat; toes 1/3 webbed; crest high trenchant and top of head a 
deep valley; brown or blackish brown with light olive, buff, 
or cinnamon area down back and a similar band or stripe on 
either side, snout 6.6-10.4 in L Nebulous toad, Bufo vallicefs 

Plate XLII 

DD. Crests absent or obscure; male excrescences on first two fingers 
not prominent, size small, below 3 in. (45 mm.). 
E. Parotoids large, low, descending on side, as long as side of 
head, body rounded; head narrower (1.7-3.15 in L.); snout 
distinctly pointed and protruding (6.9-9.1 in L.), foot with 
tarsus short (1.0-1.4 m L.), size small, 1-1 4/5 in. (16-46 
mm.); green or gray. [Recently subdivided by Taylor into 
Girard's two forms, B. debths and B. msidior.] 
(F. Toes about 1/3 webbed; nostril back from tip of snout; 
preorbital and suborbital crests smooth, without tuber- 
cles; parotoid as large as side of head; anterior edge of 
tympanum not strongly elevated. 

Little green toad, Bufo dcbtlts 
Plate XXXIV 

FF. Toes about 1/1 webbed, nostril at extreme tip of snout; 
parotoid larger than side of head; anterior edge of tympa- 
num elevated. Bufo msidtor 

Plate XXXVII 
After Smith and Taylor's 1948 checklist.) 

EE Parotoids small, raised, rounded, body flat, head broader (1.3 - 
1.6 in L.), snout not distinctly protruding, canthus rostrahs 
prominent, foot with tarsus medium (1.66-1.88 in L.), size 
medium, i 3/5-3 in. (40-74 mm ); red to gray. 

Spotted toad, Bufo punctatus 
Plate XXXIX 



Tree Frogs, HYLIDAE: Acris, Pseudacris, Hyla 

A. Alternating dark and light bands on rear of thigh; oblique white stripe 
from eye to shoulder; vertical dark and light bars on upper jaw; white 
margined triangle between eyes; hind leg very long (0.55-0.61 in L.); 
tibia very long (1.5-1.7 in L.). Cricket frog, Acris gryllus 



88 



HANDBOOK OF FROGS AND TOADS 




Figure j. Toads, Bufonidae: Bufo. i. Parotoid. 2. Tympanum (ear). 3. Warts at angle 
of mouth. 4. Gland on femur. 5. Glands on tibia. 6. Two metatarsal (sole) tubercles. 7. 
Two metacarpal (palmar) tubercles. 8. Crests united forming a prominent raised boss 
between the eyes. 9. Canthus rostralis. 10. Canthal crest, n. Preorbital crest. 12. Supra- 
orbital crest. 13. Postorbital crest. 14. Preparotoid crest. 15. Parietal crest. 16. Folds of 
skin of lower throat of male, covered at periods of rest by the lappet (17). 17. Lappet or 
apron at rear of throat of male. 18. Interparotoid interval. 




Figure 4. Tree frogs, Hylidae: Acris, Pseudacris, and Hyla. i. Tympanum (ear). 2. 
Tympanic fold. 3. Snout (muzzle). 4. Plaits on throat of male. 5. Pectoral fold across 
breast. 6. Tarsal fold. 7. Small adhesive disks. 8. Large adhesive disks. 9. Prepollex. 10. 
Rear of casque (skin fastened to skull) outlines rear of head. 



KEYS 89 

(B. Smaller; less web (3 phalanges of toe 4 free, toe i partly free); more 
rugose; anal warts less prominent; legs longer, heel reaching beyond 
snout; rear of thigh more definitely striped. Acris g. gryllus 

Plate XLV 

BB. Larger, more web (i-i^ phalanges of toe 4 free, toe i completely 
webbed); smoother; anal warts more prominent; legs shorter, heel 
not reaching to snout; rear of thigh less definitely striped. 

Acris g. crepitans 
Plate XLVI 
After Dunn.) 

AA. No alternation of dark and light bands on rear of thigh, triangle be- 
tween eyes, if present, not white-margined, no alternation of dark and 
light bars on upper jaw. 

B. No dark, brown, black, or plum-colored stripe in front of or be- 
hind eye. 

C. Head skin attached to skull; thumb rudiment apparent; rear of 
femur reticulated; no light jaw spot or dark bar between eyes; 
disks very large, size large, 2. 3/5-5 1/5 in. (64-130 mm.). 

Giant tree frog, Hyla stptentrionalis 
Plate LXXI 

CC. Head skin not grown to skull; no thumb rudiment; rear of femur 
unspotted. 

D. Rear of thigh purple; no interorbital bar between eyes; 
throat with green on either side. 

E. Green of female's throat not edged with white; back 

smooth green; body slender (head's width 3.0-3.6 in L.). 

F. Yellow or white line along side and on upper jaw; 

tibia 1.7-1.95 in L. , third finger longer (3.4-4.0 in L.). 

Green tree frog, Hyla c. cinerea 
Plate LXIV 

FF. No yellow or white line along side or on upper 
jaw, tibia 1.95-1.1 m L., third finger shorter (4.5- 
5.4 in L.). Miller's tree frog, Hyla c. evittata 

Plate LXV 

EE. Green of female's throat edged with white; back usually 
granular and dark-spotted; body stout (head's width 
X.3-2..8 in L.); white stripe from tip of snout along upper 
jaw backward. Barker, Hyla gratiosa 

Plate LXIX 



90 HANDBOOK OF FROGS AND TOADS 

DD. Rear of thigh orange or ocher; interocular bar present; throat 
without green on either side. 

E. Light stripe below eye to shoulder, back black, green, or 
brown, spotted or not; smooth; first finger 7.0-9.3 in L.; 
first toe 7-14 in L.; interorbital space 8.0-9.3 * n L. 

Squirrel tree frog, Hyla squirella 
Plate LXXII 

EE. No light stripe on upper jaw or light spot below eye, brown 
or gray usually spotted; large disks; first finger 5.1-6.5 in L.; 
first toe 5.5- 8 8 in L.; interorbital space 9.0-10.6 in L. 

Canyon tree frog, Hyla arenicolor 
Plates LX, LXI 

BB. Dark, brown, black stripe or band in front of or behind eye or both. 
C. Rear of thigh spotted. 

D. White-edged, plum-colored band from eye to groin; back green, 
unspotted with dark, rear of femur with deep orange spots; no 
interorbital bar, throat with green on either side, green white- 
edged in female. Anderson tree frog, Hyla andersonii 

Plate LIX 

DD. No plum-colored band; interocular bar present, back usually 
with spots. 

E. Network of black on yellow sides; broad dark vitta back 
from eye becoming a vertical shoulder bar, throat with 
greenish or yellow, rear of thigh netted with greenish yellow 
and purplish russet; light yellow or green spot below eye; 
size large, i 3/4-3 3/5 in. (44-89 mm.). 

Mexican tree frog, Hyla baudinii 
Plate LXIII 

EE. No network of black on yellow sides; no black or brown 
shoulder bar; throat not prominently greenish; medium to 
small. 
F. Rear of thigh brown, no netted pattern. 

G. Rear of thigh with distinct round or elliptic orange 
yellow spots; no light spot below eye, a cross-shaped 
spot on back. Pine woods tree frog, Hylafemoralis 
Plate LXVIII 

(GG. Rear of thigh specked with yellowish brown and 
darker brown; three rows of approximated spots on 
back or cruciform spot; light spot below eye present 
or absent. (After Cope.) 

Dusky tree toad, Hyla wrsicolor pbaeocrypta} 

Plate LXXVI 



KEYS gt 

FF. Rear of thigh netted with black or dark; light spot below eye. 

G. Rear of thigh with green in network; groin greenish; 

cross on back in center or rear of back; back less rough; 

intertympanic space 1.6-3.1 in L.; internasal space 8-10 

in L.; third finger 1.9-3.1 in L. 

Viosca's tree frog, Hyla aviwca 
Plate LXII 

GG. Rear of thigh with orange in network; groin orange; 
cross on forward half of the back; skin commonly rough; 
intertympanic space 3.0-3.8 in L.; internasal space o.o- 
11.5 in L.; third finger 3.1-4.8 in L. 

Common tree toad, Hyla vcrsicolor 

H. Dorsal surfaces smooth; "a number of subcircular 

golden spots in the brown ground on rear of thighs; 

interspaces [on rear of femur] often reduced to small 

circular spots." Cope's tree frog, Hyla v. chrysoscehs 

Plate LXXIV 

HH. Dorsal surfaces rough, "brown reticulation on yellow 
ground of the posterior face of the thighs"; "more 
fully marbled with yellow and brown, even covering 
the whole inner face of the tibia and the light inter- 
spaces more or less angular." 

Common tree toad, Hyla v. versicolor 

Plate LXXIII 

CC. Rear of thigh unspotted, usually a transverse bar or triangle or median 
longitudinal line between eyes. 

D. A narrow oblique cross on back; rear of thigh olive ocher or raw 
sienna; no prominent lateral dark stripe. Peeper, Hyla crucifer 

(E. Slight and indistinct chest spotting, stripe along margin of 
upper jaw, coloration less rich, dorsal stripes less broad. 

Peeper, H. c. crucifer 
Plate LXVI 

EE. More or less pronounced ventral spotting; dark spots along 
margin of upper jaw; coloration richer; broader dorsal stripes. 

Florida peeper, H. c. bartramiana Harper 
Plate LXVII 

After Harper.) 

DD. No oblique cross on back, usually a triangle or spot between eyes 
or median dark longitudinal line; dorsal and vittal stripes or rows 
of spots. 



92 HANDBOOK OF FROGS AND TOADS 

E. Size very tiny, 2.75-5/8 in. (11.5-17.5 mm.); three dorsal stripes usually 
but not always absent; stripe from eye backwards usually present; very 
long hind limbs (0.53-0.64 in L.), tibia very long (1.5-1.1 in L.). 

Little chorus frog, Pseudacris ocular is 

Plate LVI 

EE. Size small, 3/4-1 7/8 in. (19-48 mm.); 4 or 5 dark stripes or rows of 
spots usually present; hind limb 0.61-0.87 in L. 

F. Five stripes or rows of spots usually present, size small; disks in- 
conspicuous. 
Pseudacris type of coloration (middorsal, dorsolateral, and vittal 

stripes or rows of spots, 5 in number) more or less evident. 
A dark equilateral interocular triangle (sides concave or straight). 
Prominent white stripe on upper jaw, extending backward below 

prominent vittal stripe and more or less along side of body. 
A dark lateral stripe more or less evident. 
No oblique cross on back (occasionally present in Pseudacris brachy- 

phona and P. streckerf). 
Tympanum smaller, usually 11-18 in L. 
Winter or early spring breeders (December-May). 
(G. Lateral black stripe sharply defined with weak dorsal 3-stripe 
pattern; tendency for leg markings to be longitudinal; narrow 
dark line on outer tibial edge; more delicate, smoother skin; 
yellowish venter. (After Brandt and Walker )) 

Brimley's chorus frog, Pseudacns brimleyi 
Plate XLVIII 

GG. Each lateral stripe and 3 dorsal stripes more or less equally de- 
fined; leg markings more or less transverse, small size (adults 
reaching 30, 31-37 mm., 1.1-1.5 * n -)> sexually mature at 19-11 
mm. or more, Pseudacns type pattern (5 stripes or rows of spots) 
generally discernible; middorsal stripe or row of spots usually 
present; disks small, inconspicuous; body slender. 

Swamp cricket frog, Pseudacns nignta, 6 subspecies 

Plates XLIX-LIV 

H. Hind limb shorter (0.74-0.81 in L., av. 0.79); tibia shorter 
(1.36-1.8 in L., av. 1.59), foot with tarsus shorter (1.55-1.75 
in L , av. 1.69), fore limb shorter (i 06-1.11 in L., av. 1.14); 
head shorter (head to tympanum 3.19-3.63 in L., av. 3.40); 
mouth shorter (head to angle of mouth 3.73-5.71, av. 4.39). 
Minn, to B.C., Great Bear Lake, and Hudson Bay. 

Northern striped tree frog, P. n. septentrionalts 

Plate LII 



KEYS 93 

HH. Hind limb longer (0.61-0.75, avs.-o.64~o.7i), tibia longer (1.87-1.54, 
avs. 1.94-1.36); foot with tarsus longer (1.13-1.60 in L., avs. 1.36- 
1.47); fore limb longer (1.5-1.33, avs. 1.80-1.99), head longer (head 
to tympanum 1.55-3.19 in L., avs. i 94-3.14), mouth longer (head to 
angle of mouth 3.0-4.66, avs. 3.39-3.71). 

I. Tibia shorter (1.14-1.54 in L., av. 1.36); head shorter (3.14-3.19 
in L., av. 3.14); dorsum finely granular. "All the upper part an 
inconspicuous dark greenish gray with three longitudinal rows of 
large elongated spots . . ." (Wied). Middle row "unbroken ... to 
middle of the back," then "separate spots. ff N.Y. and Ont. to 
Idaho and northern Ariz. Striped tree frog, Pscudacris n. tnsmata 

Plate LIII 

II. Tibia longer (1.77-1.16 in L), head longer (1.55-3.19 in L., av. 

i 94-3-04)- 

J. Head shorter (av. 3.04 in L.), head wider (av. 3.13 in L.). "Above 
dark or fawn, with three nearly parallel stripes down the back, 
the central widening but scarcely bifurcate behind . . ." (Baird). 
Dorsum more or less smooth. N.J to N.C., possibly S.C. 

Eastern chorus frog, Pseudacris n. fcriarum 
Plate LI 

JJ. Head longer (av. 1.94 in L.); head narrower (av. 3.18-3.6 in L.). 

K. Snout shorter (5.71-70 in L, av. 633). "Above grayish 

brown or ash with distinct large circular blotches" (Baird). 

Dorsum smooth or only slightly tubercular. Texas to Kansas. 

Clarke's chorus frog, Pseudacns n. clarkii 

Plate L 

KK. Snout longer (4.6-6.66 in L., av. 5.70), dorsum more or less 
tubercular. 

L. "A light line is present along the jaw." "Above, 

speckled with small white warts, middle of back cinereous, 

with an interrupted stripe of black" (Le Conte). N.C. to 

La. Swamp cricket frog, Pseudacns n. nigrita 

Plate XLIX 

LL. "Upper lip dark plumbeous with a series of ... white 
spots" (Cope) or these absent. (Dorsum often tubercular, 
hind limb light intervals broader than in P. n. ni&nta. 
Dorsal spots smaller and better separated Brady and 
Harper). Florida. 

Florida chorus frog, Pseudacris n. wrrucosa 
Plate LIV 



94 HANDBOOK OF FROGS AND TOADS 

FF. Four stripes or rows of spots. 

G. Vittal stripe slender, dorsolateral bands curved, sometimes 
making a cross or transverse bar on back; hind limbs long 
(0.60^-0.67 in L.), triangle between eyes; disks distinct. 

Mountain chorus frog, Pseudacris bracbyphona 
Plate XLVII 

GG. Vittal mask white-bordered above; vittal stripe darker than 
other stripes or rows of spots; hind limb 0.61-0.87 in L. 
H. Body broad, toadlike (1.75-1 o in L.); disks inconspicuous; 
vitta ends at shoulder, hind limbs shorter (o 73- 0.87 in L.). 
Strecker's ornate chorus frog, Pseudacris streckeri 
Plate LVIII 

HH. Body narrow (1.94 in L.,), hind limbs longer (o 61-0.78 
in L.). 

I. Disks inconspicuous, dark mask ending beyond shoulder 
sometimes to groin, oblique groin spots light-bordered, 
interorbital space narrow (8.8-14.4 in L.). 

Ornate chorus frog, Pseudacris ornata 
Plate LVII 

II. Disks distinct, interorbital wide (7.1-10 in L ), no light 
borders around groin spots if present. 
J, Vitta may extend some distance along the side, inter- 
ocular triangle or bar absent, dark spot each eyelid 
usually present, a pair of dark longitudinal postsacral 
bars or spots the conspicuous dorsal marking, tym- 
panum smaller (14.4-10 in L.), first finger shorter 
(7.10-10 in L.), first toe shorter (7.0-13.3 in L.), 
anterior edge of tibia with heavy brown spots, lack- 
ing white line. Wrights' tree frog, Hyla wnghtorum 

Plate LXXV 

JJ. Vitta commonly ends at shoulder; interocular triangle 
usually present, no very distinct postsacral bars, the 
dorsal spotting often quite pronounced and varied, 
tympanum larger (11-14 * n L.), first finger longer 
(5.6-8.0 in L.), first toe longer (5.6-8.8 in L.). 

Pacific tree frog, Hyla regilla 
Plate LXX 

Robber Frogs, LEPTODACTYLIDAE: Leptodactylus, 
Eleutherodactylus, Syrrhophus 

A. Pupil horizontal, tympanum distinct. 

B. Fingers and toes free without distinct terminal disks, toes without 



KEYS 



95 



dermal border, size small, i 1/5-1 in. (15-49 mm.); a lateral fold; 
white band along jaw to canthus oris and humerus; dorsal color choco- 
late-brown, limbs dark cross-barred, two small tarsal tubercles; narrow 
tarsal fold, heel to orbit. White-lipped frog, Leptodactylus labtalis 

Plate LXXXII 

BB. Tips of phalanges T-shaped, toes and fingers free; terminal disks small, 
no white upper jaw stripe, no dorsolateral or lateral fold. 
C. Size large, i 7/8-3 3/5 in. (48-90 mm.), a ventral disk, voice a re- 
sounding bark, head broad (1-1.57 in L.), head wider than head to 
tympanum; eye small (7.1-9.15 in L.), eye much less than first finger; 
fingers larger, light stripe down middle of back, intertympanic fold 
present (in preserved material). 

D. Fourth toe longer; fairly closely aggregated black blotches, sides 
and hinder half of abdomen faintly areolate, skin in adults on 
dorsum stiff, coarse, areolate. Belly without vermiculations or 
spots, back covered with large black spots. 

Texas cliff frog, Eleuthcrodactylut latrans 
Plates LXXIX-LXXX 

[DD. Fourth toe shorter, upper parts either green or with scattered 
tubercles, belly with faint spots, broad light transverse band 
across back just back of fore limbs. 

Mexican cliff frog, Elcutherodactylus augusti 
Plate LXXVII] 

CC. Size small, 3/5 i 3/5 in. (15-39 5 mm), without ventral disk, voice 
a cnckethke chirp; head narrower (i 66-3.38 in L.), head usually 
narrower than head to tympanum, eye larger (5-8 in L.), eye greater 
than first finger, fingers smaller. 

D. Usually a light stripe from eye backward along dorsolateral re- 
gion, a light transverse band between eyes, toes longer, fore limb 
shorter (i 7-1.0 in L.), foot with tarsus longer. 

Ricord's frog, Elcutherodactylus rtcordtt plantrostns 
Plate LXXVIII 

DD. Usually without light dorsolateral stripe from eye backward, 
usually no transverse band between eyes; toes shorter; fore limb 
longer (i 45-1.88 in L.). 

E. Tympanum smaller (11-13 * n *-.)> hind limb shorter (0.71 - 
0.87 in L.), internasal broader (8-9 in L.); fore limb usually 
greater than foot with tarsus. 

Marnock's frog, Syrrhopbus marnockti 
Plate LXXXIV 

F. Vermiculatc dorsal pattern, wider head (ratio to L. 0.40), 
smaller size (11-18 mm.) Chisos Mts. 

Gaige's frog, Syrrhophus gaigeae 



96 HANDBOOK OF FROGS AND TOADS 

FF. Spotted dorsal pattern, narrower head (ratio of width 
of head in L. 0.36); larger size, 2.5-35 mm - Central Texas. 
Marnock's frog, Syrrhopbus mamockii 
Plate LXXXIV 

EE. Tympanum larger (8-10 in L.), hind limb longer (o-72.-o.73 
in L.), internasal narrower (9-10 in L ), fore limb usually 
less than (rarely equal to) foot with tarsus. 

Camp's frog, Syrrhophus campi 
Plate LXXXIII 



Frogs, RANIDAE: Rana 

L. Tympanum larger than eye in male, equal in female (rarely smaller in 
R. wr&attpcs), 5 .6-11.5 * n L. , throat of male differently colored, upper jaw 
unicolor, no black mask, no light jaw stripe, no regular dorsal rows of 
spots, no pronounced yellow, orange, or red on undersides of hind limbs 
or on belly or groin, tibia i 8 2. 6 in L. 

B. Rear of femur with alternations of light and dark horizontal bands; 
dorsolatcral folds absent, edge of jaw uniform. 

C. Size small i 5/8-1 5/8 in (41 66mm ), usually a light yellowish 
dorsolatcral stripe extends back from eye, two joints of fourth toe 
free of web, vocal sacs of male like a marble on either side of head, 
intcrtympanic space 3. 41- 4. 4 in L., foot with tarsus i 36-1. 56 in L. 

Sphagnum frog, Rana vtrgattpes 
Plate CXXIII 

CC. Size Urge 3 1/4 8 in. (82. 2.00 mm ), no or only one joint of fourth 
toe free of web, back uniform in color, no vocal sacs in males on 
side of head 

D. Alternation of color on rear of femur conspicuous, first linger 
generally less than second finger, mtertympanic space narrower 
(50 6 8 in L ), foot with tarsus shorter (i 4-1 67 in L ). 

Southern bullfrog, Rana grylto 
Plate XCVII 

DD Alternation of color on rear of femur not conspicuous, first 
finger generally equal to second, mtertympanic space wider 
(4.5-5.1 in L ), foot with tarsus greater (1.18-1.53 in L ). 

Bullfrog, Rana catcsbctana 
Plate XCIV 

BB. Rear of femur without alternation of light and dark horizontal bands; 
edge of jaw mottled, barred, or uniform. 

C. Dorsolateral fold absent, size large, 3 1 r-5J4 in. (81-131 mm); 
rear of femur light or white spots on brown, edge of upper jaw 



KEYS 



97 



5- 





Figttre 5. Robber frogs, Leptodactylidae: l^ptodactylus, Elcuthetodactylus, and Syr- 
rophus. i. Ventral disk. 2. Dorsolatcral fold. 3. Lateral folds. 4. Transverse (T-shaped) 
disks. 5. Subarticular tubercles sharp and saw-toothed. 6. Brachium (upper arm). 7. 
Antebrachium (forearm). 8. Femur. 9. Tibia. 10. Heel. n. Tarsus. 12. Foot. 



10 




*Figute 6. Frogs, Ranidae: Rana. T. Nostril. 2. Internasal space. 3. Costal (dorsolateral) 
fold. 4. Enlarged thumb of male. 5. Enlarged tympanum of male. 6. Tympanum of fe- 
male 7. Tympanic fold. 8. Fleshy fold on jaw. 9 Glandular folds on tibia. 10. Sacral 
hump. IT. Full webbing of male bullfrog. 12. Narrow mtcrorbital space. 13. Intertympanic 
space. 




Ft gut e 7. Narrow-mouthed toads, Brevicipitidae: Hypopachus and Microhyla. i. Broad 
waist. 2. Femur partly involved in body skin. 3. Two metatarsal tubercles. 4. Slight basal 
web. 5. Transverse fold of skin across head. 6. Depressed form of body. 7. Body thick. 
8. Legs short. 



98 HANDBOOK OF FROGS AND TOADS 

mottled; head to angle of mouth 1.0-1.7 in L.; first finger 
6.5-7.3 in L. River-swamp frog, Rana hcckscheri 

Plate XCVIII 

CC. Dorsolateral fold present, indistinct, or absent; edge of upper 
jaw uniform or barred; size medium to small; first finger 5.0-6.8 
in L.; head to angle of mouth 1.7-3. 4 * n L. 
D. Dorsal folds lacking or interrupted; back and sides mottled 
or with prominent spots, rear of femur vermiculated in its 
spotting; males with lateral external vocal sacs somewhat 
developed, size small, i 7/8-3 in. (48-76 mm.), third toe 
1.5-1.9 in L.; interorbital space 19-18 in L. 

Mink frog, Rana septtntrionalis 
Plate CXVIII 

DD. Dorsal fold on cephalic half only, cheek green with mottled 
jaw below; back uniform or with fine black speckings; rear 
of femur with fine and scant speckings; males with no lateral 
external vocal sacs; size medium, 1-4 in. (51-100 mm.); third 
toe 1.7-3.7 in L.; interorbital space 11-13 * n L. 

Green frog, Rana clamitans 
Plate CXV 

AA. Tympanum of male not enlarged (enlarged in JR. onca, rarely in R. 
artolata and R. sphtnocepbala), smaller than eye (9-19 in L.), throat not 
so differently colored in males; upper jaw with light stripe or mottling 
of light and dark, a vitta or regular spots between dorsolateral folds or 
with yellow, orange, or red on hind limbs, groin, or belly, tibia 1.4- 
1.3 in L. 

B. Dorsolateral folds absent or indistinct, low, broken; under surface 

of hind limbs yellow; skin rough, no vitta, body odor pronounced. 

C. Size larger, 1 1/3-4 1/1 in. (58-115 mm.); no outer sole (meta- 

tarsal) tubercle; no stripe on upper jaw; throat and lower jaw 

uniform or cloudy, hind limbs shorter (0.65-0 69 in L.), tibia 

shorter (1.86-1.0 in L.). Mexican frog, Rana tarabumarae 

Plate CXXII 

CC. Size smaller, i 3/5-3 3/8 in. (3 9-84 mm.); outer sole (metatarsal) 
tubercle, stripe on upper jaw present or obscure, throat and 
lower jaw spotted, hind limbs longer (0.57-0.66 in L.); tibia 
longer (1.61-1.88 in L.) Yellow-legged frog, Rana boylit 

Plates LXXXIX-XCI 

("Vomerine teeth rudimentary, on two oblique ridges between 
the nares . . . ; tympanic region not darker than rest of head; 



KEYS 



99 



fold along upper lip colored like rest of body, mottled or dark; 
red never present in coloration. Rana boylti and subspecies" 
"i. When hind leg is brought forward along the body, inside angle 
of bent tarsus and tibia reaches at least to nares and often beyond the 
end of the snout, tympanum covered with many hispid points. 
"3. A light patch on top of head, darker area crossing the posterior 
half of each upper eyelid merging insensibly into dorsal color be- 
hind; body-length under 70 millimeters. 

[California yellow-legged frog,] Rana boyln boylti Baird" 
Plate LXXXIX 

"3F. No light patch on top of head, darker areas crossing posterior 
half of each upper eyelid, when present, contrasting with dorsal 
coloration, body length reaching 81 millimeters. 

[Sierra Madrc yellow-legged frog,] Rana boyln mutcosa Camp" 
Plate XC 

"2.. When leg is brought forward, inside angle of bent tarsus seldom 
reaching beyond nares, tympanum smooth or with but a few hispid 
points; no light patch on top of head; body length reaching 73 
millimeters. 

[Sierra Nevada yellow-legged frog,] Rana boylti sierrae Camp" 
Plate XCI 

C. L Camp, Calif., 1917, p. 113.) 

BB. Dorsolatcral folds distinct full length of body (folds not conspicuous in 
R. aurora?)^ usually with a mask; skin smoother, body odor not pro- 
nounced. 

C. With mask to angle of jaw (sometimes less conspicuous or absent m 
R. aurora); males with no external vocal sacs between ear and shoul- 
der, rear of femur finely dotted and without heavy spots. 

D. No red or yellow on the under parts, mask black or dark brown, 
size small i 1/6-3 */4 * n - (19^82. mm.), tympanum larger (11- 
15 in L.) 

E. Dorsal color usually a mid-band of darker color within light 
dorsolateral folds, middorsal light stripe present or absent, 
breast more spotted. Hind limb short (0.62.^0 75 in L.), tibia 
short (i.93'-2..3 in L.); tibia usually equals foot. 

Northern wood frog, Rana sylvatica cant abri gens ts 

Plates CXX, CXXI 

EE. Dorsal color between dorsolateral fold usually like dorsum; 
no middorsal stripe; breast usually without spots; hind limb 
long (0.53-0.62. in L.); tibia long (1.6-1.88 in L.); tibia 
longer than foot. Wood frog, Rana s> sylvatica 

Plate CXIX 



ioo HANDBOOK OF FROGS AND TOADS 

DD. Some red or yellow on under parts; size average larger, i 4/5-5 in. (45- 

115 mm.); tympanum larger (11-17 * n L.) or smaller (17-11 in L.). 

E. Angle of tarsus and tibia reaching to eye, not beyond nostril; no 

mottling in groin, mask absent (if present obscure brown), inky 

spots on back sometimes light-centered, no prominent white strip 

from snout or eye to shoulder; tympanum small (16.4-11 in L.); 

eggs with i jelly envelopes, mternasal space shorter (11-16.4 * n L.)- 

F. Two metatarsal tubercles, palmar tubercles present; back and top 

of head with inky black spots, tympanum smaller (16.4-11 in 

L.); interorbital space larger (11-17.6 times in L.), fingers longer, 

especially first (6.1-6.45 in L.) and second (6.0-6.45 in L.), upper 

eyelid narrower (14.4-11 in L.). 

Western spotted frog, Rana p. prcttosa 
Plate CXVI 

FF. One metatarsal tubercle, the outer absent, palmar tubercles small 
or wanting; few irregular dorsal spots, sometimes considerable 
spotting; under parts yellow or deep orange, tympanum larger 
(13-16 times in L.); interorbital space smaller (16.1-19 i in L.); 
finger shorter, especially first (5 8-7.8 in L.) and second (5 8-7.8 
in L.), upper eyelid wider (11.6-16 in L.). 

Nevada spotted frog, Rana p. luteiventris 
Plate CXVII 

EE. Angle of tarsus and tibia reaching nares or beyond end of snout; al- 
ways or slightly mottled in groin, mask prominent black or dark 
brown (rarely, absent), prominent stripe from eye or snout to 
shoulder or shorter, tympanum normally larger (11-17 ln L.), eggs 
with 3 distinct jelly envelopes, internasal space greater (8.6 14.1 
in L.). 

F. Slight mottling in groin. Dorsum greenish brown with many 
black spots; under parts light yellow or pale salmon orange; 
third finger shorter (4 08-5.17 in L.); hind limb slightly shorter 
(0.61-0.73 in L.), fourth toe slightly shorter (1.9-1.45 m L.). 
Cascades in Washington and northern Oregon 4000 ft. or higher. 

Slater's frog, Rana cascadac 
Plate XCIII 

FF. Prominent mottling in groin; dorsum without inky or black 
spots (some few light-centered) under parts red, third finger 
longer (3.16-4.71 in L.), hind limb slightly larger (0.50-0.69 in 
L.); fourth toe slightly longer (1.51-1.15 m L.). 

Rana aurora group 
Plates LXXXVII, LXXXVIII 

G. Skin smooth; dorsolateral folds distinct; dorsum without 
spots or dotted, head to tympanum in width of head 0.93- 



KEYS ioi 

1.03, head narrower (z 5-3 .z in L.), intcrtympanic space 
(3.5-5.1 in L.); tibia shorter (1.67-1.53 in L.), first toe 
(5.4-8.0 in L.) and second toe (3.4-4.85 in L.) shorter; 
size smaller, i 3/4-3 i/z in. (44-87 inches). 

Oregon red-legged frog, Rana a. aurora 

Plate LXXXVII 

GG. Skin rough, dorsolateral folds distinct; dorsum with light- 
centered spots, head to tympanum in width of head 0.98- 
1.2.5, s * 2e larger, z 1/1-5 L /5 m - (5 8-136 mm.); head wider 
(z 15-3 o in L.), intertympanic space wider (3.0-4.45 in 
L.), tibia longer (i.46-zo6 in L.), first toe (4.5-6.5 in 
L ) and second toe (z 9-4.4 in L ) longer. California and 
Lower California. Introduced into Nevada. 

California red-legged frog, Rana a. draytomi 

Plate LXXXVIII 

CC. With no mask, males with external vocal sacs between car and shoulder. 
D. Without white line on upper jaw. 

E. Upper jaw uniform or with few dashes, size small, i 3/4-3 in. 
(44-74 mm ), dorsum unspotted or with more numerous spots 
than R. piptcns; with yellow on under parts; males with enlarged 
tympana, hind limbs shorter than in Rana onca (1.1-1.5 i* 1 L.)- 

Nevada frog, Rana fishen 
Plates XCVI, CVIII 

F. "Fewer dorsal spots and much shorter hind legs" (Linsdale). 
Hind limbs longer than R. fishert (1.46-1.8 in L.), no enlarged 
tympana. Utah frog, Rana onca 

Plates XCIX, CIX 

[The Rana fishert of Las Vegas region, Rana onca of Utah and 
Nevada, and Rana ptptens of Utah, Nevada, Arizona, and 
Imperial Valley need close examination.] 

EE. Upper jaw mottled, size larger, zJ^S^ in. (63- -113 mm.), three 
or four rows of spots between dorsolateral folds; males rarely 
with enlarged tympana. 

F. Dorsal spots large, light-centered, round with light borders, 
throat and chin clear except at sides; three or four crossbars 
on legs with small intermediates; snout shorter, head to 
tympanum z.8 to 3.z in L.; head to angle of mouth 3.15-3.8 
in L.; eye small (9.8-^.3 in L.), intertympanic space medium 
(3.8-4.7 in L.), width of head in L. narrower (z.6 to z.gz in 
L). 

(G. "Head U-shaped in outline when viewed from above; 
dorsum often smooth, or nearly so; tibia length less than 



102 HANDBOOK OF FROGS AND TOADS 

40 mm. in adults, posttympanic fold poorly developed, dorso- 
lateral folds narrow or only slightly raised or both. 

[Texas gopher frog,] liana areolata areolata Baird" 

Plate LXXXV 

GG. "Head orbiculate in outline when viewed from above; dorsum 
rugose, tibia length more than 40 mm. in adults; posttym- 
panic fold well developed; dorsolateral folds prominent. 

[Northern gopher frog,] Rana areolata ctrculosa 

Rice and Davis" 

Plate LXXXVI 
After Goin and Netting, La., 1940, p. 146.) 

FF. Dorsal spots small, irregular slightly or not light-encircled; 5 or 
8 uniform bars on hind limbs, snout longer, head to tympanum 
i. 2.^3.0 in L., head to angle of mouth 2-5-3 * i n L-> eye large 
(6.8-10.3 in L.), intertympanic space broad (^.95-3. 65 in L.); 
width of head in L. wider (i 93 - 2.. 6 in L.)> venter of head heavily 
spotted. 

(G. "Head triangular in outline, dorsolateral folds high and rela- 
tively narrow, dorsum with numerous prominent warts; 
dorsal spots poorly differentiated from gray, brown, or black 
ground color, venter always spotted at least from chin to 
midbody., dark bars on hind limbs separated by interspaces 
that are never wider than the bars. 

[Dusky gopher frog,] Kana sevosa (Goin and Netting) 

GG. "Head subtriangular in outline, dorsolateral folds low and 
very broad; dorsum smooth or lightly warty; dorsal spots 
distinct against pale ground color, chin and throat spotted; 
belly usually immaculate posteriorly, dark bars on hind 
limbs separated by light interspaces that are wider than the 
bars." Gopher frog, Ratia capito 

Plate XCII 
After Goin and Netting, La., 1940, p. 146.) 

DD. With white line on upper jaw. 

E. No orange on under parts, dorsal spots round with interspaces of 
same diameter or more. 

F. Snout shorter (60-6.8 in L), more lateral spots below dorso- 
lateral fold; tibia shorter (1.73-1.94 in L.); head to tympanum 
z.8-3.z in L.; upper eyelid wide (n.i-i4 in L.), tympanum 
normally without light center. 

Meadow frog, Kana p. pipiens 

Plate CI 

[R. pipiens complex. Plates CV-CXV.] 



KEYS 



103 



FF. Snout longer (5 .2.3-6.3 in L.); fewer lateral spots below 
dorsolatcral fold; tibia longer (1.55-1.82. in L.), head 
to tympanum 2..3&-2..S in L. , upper eyelid medium (9.3- 
11.7 in L.), tympanum usually with light center. 

Southern meadow frog, Rava p. sphenoccphala 
Plate CIV 

EE. Orange on under parts; dorsal square spots in regular rows; 
spots with interspaces less than diameter of spots. 

Pickerel frog, Raua palustris 
Plate C 

Narrow-mouthed Toads, BREVICIPITIDAE: Hypopachus, 

Microhyla 

A. Two sole (metatarsal) tubercles; basal webs on feet; snout shorter 
(9-10 in L.); usually a middorsal yellow line and mid-ventral white 
line, oblique white band from eye to shoulder. 

Taylor's toad, Hypopacbus cumus 
Plate CXXIV 

AA. One sole (metatarsal) tubercle, no webs on feet, snout longer (6.6 8.2. 
in L.), no middorsal or mid-ventral lines nor oblique postorbital band. 
B. Muzzle shorter; hind foot unusually short, hind limb short, back 
areolate, the posterior parts even pustular. 

Mitchell's narrow-mouthed toad, Mtcrobyla artolata 
BB. Muzzle longer, hind limb longer. 

C. Body depressed (thickness in length 3.0-5.6 in males, 4.0 -4.3 in 
females), tibial width in tibial length 3-4, body width in females 
2. 46- 3.1 in L.; upper eyelid 10-18 in L., eye 8.7-1^.7 in L.; skin 
usually smooth, under parts white, dorsum grayish olive. 

Texas narrow-mouthed toad, Mtcrohyla oltvacea 
Plate CXXVI 
(D. Ventrum immaculate or with scattered melanophores. 

E. A blotch or spots on the femur and tibia which form a bar 
or continuous line when the limb is folded, dorsum with 
dark spots. Taylor's Microhyla ma^atlanemts 

EE. Spots rarely present on the femur and tibia, if present usu- 
ally not forming a distinct bar when the limb is folded, 
dorsum tan and generally without markings. olivacea. 
After M. K. Hecht and B. L. Matalas, Gen., 1946, p. 7.) 
CC. Body less depressed (2-5-3.35 in males, z. 65-3. 4 in females), 
tibial width in tibial length 2-5-3 - times, body width in females 
1.83-1.3 in L., upper eyelid i8.6-z8 m L., eye icx-i4 m L., skin 
smooth, tuberculate, or pustular; under parts gray, or brown 
speckled and mottled; dorsum black, gray, or brown. 

Narrow-mouthed toad, Microhyla carolinensis 
Plate CXXV 



Accounts of Species 

ORDER SALIENTIA Laurenti 
FAMILY ASCAPHIDAE 

Genus ASCAPHUS Stcjneger 

Map 4 



THE first recognition of this form by Dr. Stejneger and the subsequent 
studies of it by Dr. Helen Thompson Gaige were two of the outstanding 
events in North American batrachology in the last forty-five years. This 
"tailed" frog has relatively the broadest internasal space of all our frogs, 4.66 
to 8 into length, whereas all others range from 8 (or 7.77) to 21.4 into length. In 
hind limb it is relatively longer than robber frogs, narrow-mouths, spadefoots, 
or toads and comparable to tree frogs and frogs. It has long fore limbs. Pos- 
sibly both fore and hind limbs are long for climbing. Of the first live ones we 
had, all except one escaped by crawling up the inner sides of a barrel. It has 
a large mouth, snout, eye, interorbital space, and upper eyelid. 

The relationships of this frog have been much discussed. When Stejneger 
(1899) with rare courage and fine discrimination described this form, he 
called it a Discoglossid. Later, VanDenburgh (1912) found it to be not "ecau- 
date," and followed Stejneger. American authors such as Barbour and Coch- 
ran (1932) and most foreign workers like Boulenger (1910), Gadow (1920), 
Nieden (1923), and Perrier (1925) kept it within the Discoglossidae. From 
1924 to 1931 and until his death Noble placed it in the Liopelmidae, and Vil- 
liers (1934) and Pusey (1943) are in agreement with this view. In 1923 Fejer- 
vary created a special family, the Ascaphidae, and in the third to fifth editions 
of Stejneger and Harbour's check list (1933, 1939, and 1943) these authors 
adopt this family name, which we now follow. We have seen considerable 
material both preserved and alive and are much impressed with its Discoglos- 
sid affinities. 



RIBBED TOADS: ASCAPHIDAE 105 

American Bell Toad, American Ribbed Toad 

Ascaphus truet Stejneger. Plate XVI; Map 4. 

Range: Washington, Oregon, northern California (south to Dyerville; 
Myers, 1931, 1943), Emida, Idaho (Ehrhardt, 1946), western Montana (An- 
derson and Smith, Hazzard, 1932; Donaldson, 1934; Rodgers and Jellison, 
1942; Shpp and Carl, 1943), British Columbia (Ricker and Logier, 1935; Slipp 
and Carl, 1943; Carl, 1945). 




Map 4 

Habitat: These "toads" live in forested sections, under rocks in perennial, 
usually swift-flowing, small mountain streams of low temperature. After 
heavy rains several collectors have found them in moist woods, at varying dis- 
tances from the streams. 

Size: Adults, i %-2 inches. Males, length to tail 29-42 mm., tail 3-10 mm. 
Females, 28-51 mm. 

General appearance: This small "toad" is gray, pink, or brown to almost 
black. Numerous black spots occur on the tops of the legs and on the back. 
The bar across the head is pale green to pale yellow; the underside of legs 
rose. The females are usually lighter than the males. The brown parotoid 
gland is well developed or may be broken into a glandular ridge along the 



io6 HANDBOOK OF FROGS AND TOADS 

side. The fingers are long and slender, free of webbing; the toes, slightly 
webbed. The head is flattened, slightly broader than long, the snout obtusely 
pointed, and with no visible tympanum. The skin is smooth or slightly rough- 
ened with granules, wrinkles, warts, and small tubercles. The conspicuous 
"tail" of the male is level with the ventral side of the body. This is %-% inch 
(3-10 mm.) long, %-% inch (4-5.5 mm.) broad. The anus is a large swollen 
orifice just ahead of the constricted tip. The female appears much like a tree 
toad. 

Color: Carbon River Valley, Rainier National Park, from J. R. Slater, May 
16, 17, 1930. Female. Middle of back from eye bar to rump grayish olive or 
citrine drab. Parotoid wood brown, cinnamon, or mikado brown. From paro- 
toid along side is pecan brown, mikado brown, orange-cinnamon, or sayaFs 
brown. Same wash on upper parts of tibia and an area near anus. Vertical part 
of snout except central bar (like bar between eyes) black, aniline black, or dull 
purplish black. Area below eye ageratum violet or vinaceous-purple. Bar from 
snout to eye and behind eye to shoulder insertion same color as vertical snout. 
On top of forelegs and hind legs and along side a few larger black spots. Along 
sides these arc more elongate below, defining the browns. Color of hind legs 
and forelegs same as mid-back. Bar across head at front of eyes glass green or 
deep seafoam green. Same color on side of face below black nostril, eye, and 
shoulder insertion bar. This bar bordered above with same color. Top of snout 
mignonette green or ecru-olive. Underside of body except belly and femur 
dull citrine, ecru-olive, or barium yellow with interstices of vinaceous, dark 
anthracene violet or dark maroon purple. Some of belly light orange-yellow 
with some vinaceous or pinkish vinaceous interspersed. Femur pure deep 
vinaceous or purplish vinaceous. Ins: bar across top of vertical pupil cream 
color or pale chalcedony yellow with some reticulum of burnt sienna or hazel. 
This color in upper part of eye obscures the cream or yellowish. Lower eye 
auburn, chestnut, or chestnut-brown with little specks of orange-rufous. 

Male. Back kaiser brown, hazel, tawny, cacao brown, or vinaceous-russet. 
Top of hind legs and forelegs with numerous black spots, so also back; bar 
from orbit to orbit pale chalcedony yellow. Top of snout deep olive-buff. Ver- 
tical snout bar like eye bar. Male has venter, all except femur, like female de- 
scribed. Two patches on arm and base of finger black or seal brown or warm 
blackish brown. Iris: see female. 

Structure: The tail is an intromittent organ; lower jaw from below an al- 
most perfect semicircle; second, third, and fourth vertebrae bearing short ribs; 
tongue attached by broad surface, and cannot be protruded; arms of male con- 
spicuously heavier than in the female, as also are the hind limbs and feet. 
Breeding males with dark excrescences on the inner edges of the inner fingers 
and along the inner edge of the forearm, a ball-like excrescence at the base of 
the thumb, and often dark excrescences on each side of the breast; tuberculcs 



RIBBED TOADS: ASCAPHIDAE 



107 



much finer on the rear portion of the back and on the hind limbs in the fe- 
male; eye large with a vertical pupil. 

Voice: No vocal sac. 

Breeding: Known dates are in May, June, July, August, and September. 
The eggs in circular masses of rosarylike strings are attached to the underside 
of stones in creeks. The eggs are not pigmented, few in number, and very 
large, % inch (8 mm.), the yolk % inch (5 mm.) in diameter. The mature 
tadpole is medium in size, i%-2%c inches (42-55 mm.), its body round, its 
tail long, its crests not conspicuous. The tooth ridges are %o (upper ridges 
with more than one row of teeth). The period of development is probably 
over winter. They transform during July, August, and September at %-% 
inch (14-18 mm., possibly 20 mm.). Wallace Wood (Calif., 1939, p. no) found 
tadpoles in January. Ricker and Logicr (B.C., 1935, p. 46) in their five British 
Columbia streams found one-year tadpoles 29-41 mm. and a second-year tad- 
pole 42 mm. 

Journal notes: March 16, 1942, Russ Grove, northern Humboldt Co., Calif. 
Mr. Milne says Klamath River is the line of separation for many forms such 
as Ascaphus. 

March 17. Went to south side of Wilson Creek, Del Norte Co., Calif., 8 miles 
north of Klamath and close to the ocean, according to directions of Wallace 
F. Wood. With seine and hoe we got plenty of Ascaphus tadpoles and larvae 
of Rana boylii boyht and Dicamptodon ensattis. There were no adult frogs or 
salamanders under stones m creek or under outlying ones or boards. When 
you pick them up, the Ascaphus larvae attach themselves to the finger or net. 
Caught against the side of the can, the yellowish white tip with black crescent 
band behind it is conspicuous. 

March 20. Went up MacKenzie River watershed with Ruth E. Hopson of 
Springfield, Ore. . . . At last caught one Ascaphus trnei tadpole. 

March 23. With Kenneth Gordon up Santiam River. . . . Sixteen and one- 
half miles from Foster at mouth of Moose Creek. Storm, Livczey, and I 
pulled seine in creek for Ascaphus. Didn't get any, but they are here. An old 
student, Dr. A. C. Chandler, secured them in Santiam National Forest 25 
years ago. 

March 29. Near Lake Cushman dam; couldn't get very close because of war 
regulations. Made a long loop back to the highway through the lumbered 
countryside. Oh, the desolation of that Hamma Hamma River country! Dr. 
H. T. Gaige's historic collecting site is under many feet of water. 

March 31. With J. R. Slater. Went in afternoon to Carbon River just outside 
Rainier Park entrance. In a tumbling hillside cataract we saw little A. truei 
tadpoles attached to rocks; their tails were wagging in the current. Caught no 
adults. . . . Drove up to ranger cabin but soon came back and tried another 
little stream which also has Ascaphus tadpoles. 



io8 HANDBOOK OF FROGS AND TOADS 

Authorities' corner: Mrs. H. T. Gaige (Wash., 1920, pp. 2-3), working in 
the Olympic region of Washington wrote: "It was under the rocks in these 
little creeks that Ascaphus lived. . . . One found them only by working 
slowly upstream and turning over every movable stone. Usually they either 
floated down the stream with the debris which was released when the stone 
under which they were resting was overturned, in which case it took quick 
action to catch them before they were out of sight, or they made no effort to 
move, and in the shifting lights and shadows their color so closely resembled 
a bit of fir bark or the small red stones which were abundant in the streams 
that they were distinguished with difficulty. Occasionally they were alert and 
slipped away like a shadow. When placed on land they were awkward and 
stupid in action and appearance and made little effort to escape. They were 
solitary; never more than one was found under a single stone and individuals 
were usually well separated in the stream." 

"The temperature of the water at i P.M. registered 10 Centigrade. Here 
the tadpoles were very abundant, for as many as seven were seen and collected 
from an area that could be covered by the palm of one's hand. They were first 
observed in the clear water attached to the rocks on the bottom of the creek. 
Efforts made to capture them caused them to release their holds and swim up- 
stream for a few inches and there seek shelter under the rocks. Here they were 
more difficult to see since their coloration blended nicely with the debris under 
the rocks, but once one knew where to look and what to look for, they were 
more obvious" (A. and R. D. Svihla, Wash., 1933, p. 38). 

"Trip July 14-15, 1941. The tadpoles were at approximately the same stage 
of development as those taken on June 22, 1930 (which are now in the U.S. 
N. M.). They have short hind legs. The black caudal area and milk-white tail- 
tip are prominent in most of the individuals. 

"I watched one tadpole feeding on a stone over which the swiftest current 
in a riffle was falling. He moved about readily when he desired, a millimeter 
or two at a time, by a slight 'inching along' movement of the adhesive mouth- 
parts. He would turn this way and that, his tail fluttering and flapping down- 
stream in the current, and the slight but constant movement of his mouthparts 
indicated that he was scraping at the surface of the rock. Although nothing 
could be seen on the stone, there was probably a growth of minute algae. At 
times the tadpole was almost half out of water, and again he would browse 
along till he was nearly on the underside of the stone. All of the tadpoles seen 
were in the swiftest current available, usually on the side of a stone where a 
drop of a couple of inches formed a riffle" (G. S. Myers, Calif., 1943, p. 126). 

"Herewith are being sent to you specimens of the larvae of Ascaphus truei 
that I collected during the past summer. These specimens were collected July 
30, 1946 near Emida, Idaho. They are from Township 43 North, Range 2 
West, Section 13, East Fork of Charley Creek. 

"This area is mostly in the classification of open reproduction, having been 



RIBBED TOADS: ASCAPHIDAE 



109 



burned several times in the past. The streams have numerous beaver dams. 
These particular specimens were gathered where the water was free flowing 
and the stream bed of gravel. They were common lying on the top of the 
gravel and offered no resistance to capture. I was unable to locate any adults. 
". . . The boys in the camps at first enjoyed swimming in one or another 
of the beaver ponds, but they soon began to complain of the 'leeches' which 
attached themselves to their legs and toes. I found that these were the same 
larvae. The ponds accumulated a silt deposit but were still water" (letter from 
R. P. Ehrhardt, Kenyon College, Ohio, Sept. 6, 1946). 



no 



HANDBOOK OF FROGS AND TOADS 





Plate XVI. Ascaphus truei (XO- i>5- Fe- 
males. 2. Tadpole. 3,4. Males. 



Plate XVII. Scaphiopus couchii. 1,6. 
Males (X%). 2. Male (Xtt). 3- Female 
(X%). 4- Male croaking (X%). 5- 



FAMILY SCAPHIOPODIDAE 
Genus SCAPHIOPUS Holbrook 

Maps 5-7 

Couch's Spadefoot, Southern Spadefoot, Rain Toad, Sonoran Spadefoot, 
Sonora Spadefoot, Cape St. Lucas Spadefoot 

Scaphioptts coitchii Baird. Plate XVII; Map 5. 

Range: Texas to Arizona and Utah, northern Mexico and Lower California. 

Habitat: These spadcfoots live in subterranean burrows, often under logs or 
similar shelter, and are nocturnal in habit. They breed in temporary pools, 
coming out only after heavy rains. 

Size: Adults, 1%-$% inches. Males, 48-70 mm. Females, 50-80 mm. 

General appearance: The short, fat, toadlike body has a greenish back, more 
or less marbled. The skin is roughly tuberculate, with many light tubercles on 
the sides. Both the ear and parotoid glands are indistinct. The eyes are large 
and protuberant with vertical pupils. The venter is whitish; the fingers and 
toes are light; and the rear of the arm and leg has a light band. The outer sole 
tubercle and sometimes the tips of fingers and toes are dark. 

Color: Male. Beeville, Tex., March 26, 1925. Upper parts yellowish oil green; 
on sides same, oil yellow mixed with yellowish oil green; tubercles on side, 
posterior back, and tibia calliste green; tubercles below vent white; rear of legs 
and groin parrot green; below eye and lower muzzle buffy citrine; tympanum 
oil yellow or yellowish oil green and a little livid brown. Under parts white 
except small pectoral patch of Martius yellow or pale greenish yellow, and area 
from groin to groin across rear of belly where only livid brown or purplish 
vinaceous exists. Eye: pupil vertical, rim lemon yellow, iris lemon yellow to 
bright green-yellow reticulated with black. 

Female. Comfort, Tex., May 19, 1925. Bands on front and rear of upper 
eyelid citron green or oil yellow; same color on side of upper jaw, above and 
below tympanum, along sidfcs, on back, on dorsum of hind limbs, in each 
case interspersed with oil green. These two colors are sharply contrasted in the 
female, while in the male the color is a more uniform calla green or oil green, 
with little spotting of lighter color. Black bars on the limbs, and some black 
on nostril. Another has mignonette green on sides spotted with black and ivy 
green, tea green, or water green on back. Fingers white on dorsum, toes with 



112 



HANDBOOK OF FROGS AND TOADS 



some white on clorsum. Outer bands on feet whitish. Under parts white except 
lower belly and under legs, which are slate violet (2) to vinaceous-lilac. Iris 
green-yellow or bright green-yellow with black on outside, which is sometimes 
a horizontal bar in front and behind. From black on outside, black lines run 
almost to the vertical pupil. In general one can identify females because they 
are spotted, while males are more uniform. 

Structure: No pectoral gland; skin on crown of head thin; tongue subcir- 
cular, slightly emarginate; hand nearly as long as forearm; toes fully webbed. 

Voice: The chorus is harsh and noisy, a great caterwauling, and can be heard 
a considerable distance. The individual call tvow, me ow, or a ow is a most 
unearthly noise like that of someone in pain. 




Map 5 

"The cry of the male is a loud, resonant 'ye-ow' repeated at intervals; that 
of the female, a short grunting 'ow 1 uttered several times in shorter intermis- 
sions 11 (J. K. Strecker, Jr., Tex., igoSe, p. 203). 

"In the case of S. cottchii it [the song) is a bleat lasting from 5-6 seconds 
ordinarily. This can be best described as sounding so much like the bleat of 
a lamb as to be mistaken for it. The song is given while the male sits upon the 
mud near the edge of the puddle" (A. I. Ortenburger, Ariz., 1925, p. 19). 

Breeding: The time is from April to August at periods of heavy rainfall. 
The eggs are in bands Vi inch (6 mm.) across or cylindrical masses on plant 
stems, the jelly is rather firm, the eggs are close together, black above and 
creamy white below. The vitellus is Ms-He inch (1.4-1.6 mm.). In warm 
spots they hatch in iH days. The "bronzy" tadpole is black, dotted with old 



SPADEFOOTS: SCAPHIOPODIDAE 113 

gold or fawn, small, i inch (24.5 mm.), broad with tail tip rounded. The 
tooth ridges are %, rarely %, %, %, %. After a period of 15-40 days, the tad- 
poles transform during the summer months and early fall at %o-% inch (7.5- 
12.5 mm.). 

Journal notes: May 29, 1925, Comfort, Tex. The spadefoots are calling as 
they float spread out on the surface. Their sides are swelled out and vibrating. 
They often seem to curve their backs in their tremendous efforts. 

June 4. In Comfort, Tex., in roadside pools where, on May 29, we found 
Couch's spadefoot breeding, there are now large tadpoles. The rain came May 
28. The eggs must have been laid then. How fast! 

June 7. At Encinal, Tex., roadside pool alive with spadefoot tads. . . . 
More spadefoots 2 miles beyond Cactus. Whenever the pools begin to dry up, 
grackles go there to eat the tads. 

June 8. Near Dolores, Tex., we stopped beside a long roadside rain pool. It 
is a sandy area with scattered bushes, very little herbaceous material, and mes- 
quite rather far apart. All stages were here, even tiny transformed ones leaving 
the pond. They were hopping out so thickly that they formed a seething mass, 
several spadefoots deep. They were gathered around small herbs, small bushes, 
and larger ones, when possible, for shade, and generally just enough in a place 
to match the shadow of the plant. Many of them crawl out of the pond when 
they still have very long tails. I wonder if this hastens the shrinking of the tail. 

June IT, east of Hebronville, Tex. Last night we went out with flashlights. 
Near a clump of cactus saw a frog. It proved to be Couch's spadefoot. Caught 
it. The instant we released it, it started for a clump of prickly pear and went 
down into a pack rat's hole. I verily believe this is one place where they aesti- 
vatc. 

Authorities 9 corner: 

J. K. Strecker, Jr., Tex., 19086, p. 203. F. W. King, An/,., 1932, p. 175. 
A. I. Ortenburger, Ariz., 1925, p. 20. L. W. Arnold, Ariz., 1943, p. 128. 

Hammond's Spadefoot, Western Spadefoot, Hammond's Spea, Western 
Spadefoot Toad, New Mexican Spea 

Scaphiopus hammondn hammondn Baird. Plate XVIII; Map 6. 

Range: California to Lower California, cast through Arizona, southwest 
Colorado, New Mexico, to Texas and Mexico. 

Habitat: They live underground in burrows which they dig in soft earth 
by backing into the ground and digging with their hind feet, which are armed 
with spades. They rock the body as they dig, and the dirt falls into the bur- 
rows on top of the toads. They breed commonly in temporary rain pools or 
temporary overflow areas. 

Size: Adults, i%-2% inches. Males, 37.5-59 mm. Females, 37.5-61 mm. 

General appearance: The body is stout and toadhke, small in size. The eyes 



HANDBOOK OF FROGS AND TOADS 



are large and protuberant with vertical pupils. The skin is fine, relatively 
smooth, dotted with fine roundish tubercles. The back is greenish, the sides 
yellowish glaucous, light mineral gray, or greenish. There are green spots on 
the back, top of head, and legs. The forward under parts are white, sometimes 



M- 

: '.V.V IT - 




Map 6 

buffy on the throat; the rear under parts purplish. The males have a wash of 
grayish green on each side of the throat. 

Color: "Color (in life) above dark green, with scattered spots of dusky; four 
(or two) incomplete longitudinal stripes of dull white, inner pair in line with 
inner margins or orbits, the outer in line with tops of tympanic membranes; 
tubercles on back and sides tipped with red or orange in young individuals; 



SPADEFOOTS: SCAPHIOPODIDAE 115 

ventral surface plain white; throat region (vocal sac) blackish in males, dusky 
in females. Females lack the conspicuous longitudinal dorsal streaking of 
white, this being broken and much less extensive" (T. I. Storcr, Calif., 1925, 
p. 149). 

Structure: Head broader than long, muzzle short and overhanging the 
lower jaw; epidermis on top of head thick and horny; tympanum indistinct; 
no parotoid gland; tongue very large, entire; no tibial or pectoral gland; hind 
limb, tibia, foot, fourth toe, and fingers relatively longer than in Couch's or 
in Holbrook's spadefoot. 

Voice: The males call, lying on the surface of the water. The call is a rolling 
or bubbling one, a croak more like the croak of some frog than Couch's or the 
hermit spadefoot. It has been described in widely different ways: as the loud 
purr of a cat with the metallic sound of grinding gears, as a low-toned ttrr-r-r-r, 
as a loud crah-crah-t ah, and as a resonant ye-ow. It has been called unusual, 
weird, plaintive, and ventriloqual. 

Breeding: They breed from mid-February to August, dependent upon 
heavy rainfall. The eggs are in cylindrical masses attached to grass or plant 
stems. The eggs on the periphery of a jelly cylinder may look stalked, the stalks 
%-% inch (5 or 6 to 9 mm.) long and Vio-/io inch (1.4-2.3 mm.) in diame- 
ter, the eggs Msn-Mo inch (1.0-1.6 mm.). The eggs hatch in i%-2 days. The 
dark greenish black tadpoles may grow large, 2% -2% inches (65-70 mm.) 
long. They are broad, almost round-bodied in dorsal view, the eyes close to- 
gether, the tail short with rounded tip, the spiracle low, almost ventral. Like 
most spadefoot tadpoles, the musculature of the tail stands out very promi- 
nently. The tooth ridges are %, %, %, %> % After 30-40 days the tadpoles 
transform from May 20 to September i, at M>-i!4 inch (13-32 mm.). The 
tadpole is carnivorous in habit and may prey on its own kind, but it is a very 
effective enemy of the mosquito. 

Ortenburger (Ariz., 1925, p. 19) wrote: "(a) Diameter of egg including 
gelatinous layers S. couchit 2.5-3.5 mm.; S. hammondn, 1.5-2.0 mm.; 
(b) thickness of jelly layers S. conchii, usually more than i mm.; S. ham- 
mondit, mostly less than 0.5 mm.; (c) method of attaching the individual eggs 
to others in mass S. cotichh attached with very short stalk and S. hammondn 
by a slender stalk 5-10 mm. in length . . . ; the eggs of S. hammondti were 
arranged on similar objects but arranged spirally around them, thus differing 
from S. cottchii in which no spiral arrangement could be easily made out." 

Our measurements closely follow those of Ortenburger. The egg masses on 
stems may be from 6 or 7 to 25 mm. in diameter. Egg stalks may be from 5-9 
mm. long and 1.4-2.3 mm. in diameter; vitellus 1.0-1.6 mm.; hatching period 
i%-2days. 

Journal notes: July 8, 1917. Quite a rain fell near Sierra Blanca, Tex. At 7 
o'clock, we heard no notes in the creek, but later from our camp one-half mile 
away, we heard the chorus plainly and decided it must be spadefoots. We 



ii6 HANDBOOK OF FROGS AND TOADS 

found toads and spadefoots of two species migrating from the mountain side 
of Sierra Blanca toward the pool and noise. . . . Along the edges of the swift 
stream now flowing across the flooded area we found Scaphioptts couchii. 
Their cries were catlike. The 5. hammondii were on the surface of the water 
and their calls were bubbling. . . . The Hammond's male will float like 
S. holbroofyi. When he croaks, the rear half of the back dips beneath the 
water. 

July 9. The stream has disappeared; it is now broken up by intermediate 
mud flats. The spadefoots and toads have disappeared from last night's ren- 
dezvous. 

July 27, 1925. Found Hammond's spadefoot eggs, tads, and transformed in 
ponds by the roadside 16 miles east of Vail, Ariz. In one pond must have been 
40-50 masses of eggs on the ends of grass stems. The eggs seemed to be out 
on periphery of the jelly cylinder. They look to be stalked. This is the same 
thing we found in Texas Pass 8 years ago, when we suspected it might be 
Hyla arenicolor, but now we know it is S. hammondii. These eggs were laid 
last night and are already nearing hatching. 

April 20, 1942. With Hadsell and Culbertson to Fresno, Calif., slough. Be- 
fore we reached White Bridge we came to two or three ponds where we took 
large light-colored tadpoles (S. h. hammondii). Here Culbertson took some 
that were almost transformed, earlier. 

June 13, Mesa, Ariz. Went out last night toward Florence Junction. Heard 
bullfrogs and Hammond's spadefoot in a pool at Desert Wells. 

July 9, Lakeside, Ariz. Tonight, early in the evening heard no frogs. After 
I went to bed about 10 P.M. heard a few S. h. hammondii in one of the ponds 
above us. 

July 10. Tonight, heard a few isolated S. h. hammondii again. 

Authorities' corner: 

E. D. Cope, N. Mex., 1884, p. 14, 

T. I. Storer, Calif., 1925, pp. 156-157. 

F. W. King, Ariz., 1932, p. 175. 

Central Plains Spadefoot, Central Plains Spadefoot Toad, Cope's Spea, 
Western Spadefoot Toad 

Scaphtopus hammondii bombijrons (Cope). Plate XIX; Map 6. 

Range: Not fully defined. Stejneger and Barbour apparently consider the 
form from the Dakotas to Oklahoma, northwestern Texas, New Mexico, and 
west to Idaho to be S. h. bombifrons, but V. M. Tanner (1939) considers those 
west of Utah's east line, north of northern Arizona, and east of Nevada's west 
line to be Cope's old form inter montanus. H. M. Smith (Kansas, 1934) re- 
marks that the range overlaps with S. h. hammondii in Colorado, western 



SPADEFOOTS: SCAPHIOPODIDAE 



117 





Plate XVIII. Scaphiopus hammondii 
hammondii (X%)- i>M> 6 * Males - 3- 
pole. 5. Transformed frog. 7. Eggs. 



Plate XIX. Scap/nopus hammondii bom- 
bifrons (X%)- 1,2,3,5.' Females. 4. Male. 



n8 HANDBOOK OP FROGS AND TOADS 

Oklahoma, and northern and western New Mexico. P. Anderson (Mo,, 1945) 
recently added Missouri to its range. 

Habitat: "East of Colorado Springs in the low rolling hills is one of these 
areas. The soil is a mixture of sand, gravel, and loam and generally quite dry. 
It is here that adult spadefoot toads are found at depths varying from a few 
inches to several feet. ... It usually chooses soft ground in which to burrow. 
With its spade-armed feet it pushes the soil aside, and by a slow rocking move- 
ment sinks backwards beneath the surface of the ground. The heavy skin of 
the head is probably used to keep the burrow open in front or to pack the 
earth of the walls of the burrow. The descending toad leaves no trace on the 
surface to indicate its course" (R. J. Gilmore, Colo., 1924, pp. 1-2). 

"Along the Powder River near Powderville in Montana, on June 15, 1916, 
while lying upon my cot, I heard a curious rustling in the dry leaves about our 
tent. Upon investigation with a flashlight many small spadefoot toads were 
found. They were hopping about in the dry leaves which were scattered about 
on the sandy soil. When hunted with a flashlight they endeavored to burrow 
out of sight and but a few minutes were required for them to entirely conceal 
themselves. These spadefoots make circular holes in the ground and yet in 
sandy soil it is very difficult to find the place where they have burrowed down, 
for in most cases it seems as if they had pulled the hole in after them. After 
the breeding season is over they take more pains in constructing their burrows, 
as ... [the burrows] are well rounded and resemble somewhat an earthen 
jar with a narrow top. Around this opening there is present some sticky mat- 
ter which may aid in the ensnaring of insects. I have usually found this toad 
most plentiful in sandy areas, especially along the banks of streams, though 
they occur on the elevated plains from Kansas to Montana" (R. Kellogg, 
Mont., 1932, p. 36). 

Size: Adults (roughly) i%-2% inches. Males 38-52 mm. Females 40-57 
mm. 

General appearance: Stout; small; vertical pupil; interorbital boss. Grayish, 
reticulated or uniform; white below except throat of male. Two series of dorsal 
stripes or these absent. Smoother than S. hammondii. Internasal distance i% 
in eye to nares in bombtfrons, i in hammondii. 

Color: Colorado Springs, Colo., July, 1928, from R. J. Gilmore. Female. 
Sides yellowish glaucous or light mineral gray or water green; back in gen- 
eral, top of head, and hind legs vetiver green, tea green, or water green. Back 
of each eye on back is a dull citrine, water green, or yellowish glaucous spot 
edged with Lincoln green or deep grape green. On hind legs, rear of back, and 
top of head are spots Lincoln green or dusky olive-green. Some of these spots 
have light centers which are tubercles (under the lens, dark olive-buff in 
color). Under parts are white or cartridge buff, particularly on the throat; 
other parts are congo pink to salmon-buff or light ochraceous buff. There are 
two tubercles ahead of vent and also beside it that are white to light buff. Iris 



SPADEFOOTS: SCAPHIOPODIDAE 119 

pale greenish yellow or light green-yellow unbroken around pupil; rest with 
black lines. Iris may be orange-pink. A spot or transverse bar like dorsal spots 
occurs on upper eyelid. 

Male. The males on either side of throat have wash of deep bluish gray- 
green, or light terre verte, or grayish blue-green. [This from memory of males 
alive three days ago.] In alcohol the throats of male are very bluish or plum- 
beous. 

Structure: Metatarsal tubercle rounded; parotoid gland absent. "A Scaphio- 
pus with rounded, not elongated, inner metatarsal tubercles; tip of fifth toe 
frequently blackened and corneous; anterior interorbital region swollen, con- 
vex; parotoid glands indistinct, tympanum also usually; toes nearly fully 
webbed, fingers very slightly; pupil vertical" (H. M. Smith, Kan., 1934, 
p. 427). 

Voice: "The call of this toad is quite weird and unusual and may be likened 
to the squawk of some animal when severely injured or a resonant ye-ow. 
Once heard this distinct call is not likely to be forgotten" (R. Kellogg, Mont., 
1932, p. 36). "After arriving at the ponds the male spadefoot indulges in very 
vigorous nuptial song, which continues without interruption until the mating 
has been completed. The efTect has been described as 'weird plaintive cries,' 
'hoarse and woeful' " (R. J. Gilmore, Colo., 1924, p. 3). 

Breeding: These frogs breed from May to August in the rainy season. The 
amplexus is inguinal. "The egg masses vary in size. Large masses contain 200 
to 250 eggs, smaller ones 10 to 50. The mass is attached to submerged vegeta- 
tion, or to any object protruding from the bottom. The mass is elliptical in 
shape." "The incubation period as observed in the field seems to be less than 
forty-eight hours." "Two and one-half inches is the maximum length of the 
majority of adults in any tadpole community." "In 1921, specimens were 
found completely transformed after thirty-six to forty days" (R. J. Gilmore, 
Colo., 1924, pp. 4, 5). The tadpole is carnivorous and herbivorous. 

From conversations, letters, material, and experience, it is evident that our 
friends, Gilmore and Kellogg, know this form best. Dr. Gilmore has used this 
species in classes for a quarter-century and Dr. Kellogg has known it equally 
long. In recent years A. H. and M. S. Trowbridge and A. N. Bragg of Okla- 
homa have added the most to our knowledge of this frog. 

In 1934 H. M. Smith characterized the tadpole thus: "Upper mandible with 
a large median beaklike projection, lower mandible with a deep elevated me- 
dian notch; a black, corneous toothlike projection from roof of mouth; buccal 
musculature conspicuous and visible through skin as are viscera!" In 1929 we 
had described the tadpole of S. hammondii with no thought of separation into 
its three forms. 

In 1942 A. N. Bragg (Okla.) characterized the tadpole of S. h. bombifrons 
as having "jaws without a beak and notch; jaw muscles not overdeveloped." 
In 1941 in New Mexico he discovered that "Smith's figure and description are 



120 



HANDBOOK OF FROGS AND TOADS 



of S. hammondii, and R. J. Gilmore (1924) and Wright's (1929) are of S. 
bombifrons, although labeled the reverse of this." 

Authorities' corner: 

J. A. Tihen and J. M. Sprague, Kans., 1939, pp. 501-502. 
G. A. Moore and C. C. Rigney, Okla., 1942, p. 78. 

Great Basin Spadetoot Toad, Western Spadefoot Toad 

Scaphiopus hammondii intcrmontanus (Cope). Plate XX; Map 6. 

Range: Utah, Nevada, northern Arizona, western Colorado, southwestern 
Wyoming, Idaho except extreme north, southwestern Washington, eastern 
Oregon a Great Basin form (after V. M. Tanner). British Columbia. 

Habitat: Canyon pools; desert springs and pools, intermittent and perma- 
nent; irrigation ditches; stream edges; rain puddles; water pockets; water de- 
pressions made by cattle. 

Size: Adults iM-2% inches (40-63 mm.). Males, 40-59 mm. Females, 45- 
63 mm. 

General appearance: "The evolution of the subgenus Spea seems to be from 
hammondii through bombifrons to tntermontanus. In these species there is a 
progressive development of the osseous parts of the cranium with a closure 
of the fronto-parietal fontanelle in practically all specimens of intermontanus" 
". . . Fairly rugose." "Intermontanus has a greater internarial distance than 
either bombifrons or hammondii" (V. M. Tanner, Gen., 1939, pp. 15, 16). 

"They appear to demonstrate that bombifrons and hammondii are in the 
same species, for there is obvious intergradation in every character. Moreover, 
it seems likely that additional material from this intervening area [Nevada] 







Plate XX. Scaphtopus hammondn tnterwontanus (X%)- From C. L. Patch, 
Okanagan Landing, B.C. Adults. 



SPADEFOOTS: SCAPHIOPODIDAE 121 

would demonstrate the validity of a third race, intermontanus" "Compared 
with hammondii this form was supposed to be distinguished by larger size, 
lighter colors, and the presence of the superior pair of light lines" (J. M. Lins- 
dale, Nev., 1940, p. 200) . 

Color: "Their color was yellowish olive above, spotted with darker olive, 
belly soiled white, chin darker, sides of front legs and feet gray, the hind legs 
tinted on the under surface with blood red. The eye was large and very bril- 
liant, the iris brassy with fine black reticulations. The pupil was very sensitive 
to light. There were two broad, grayish stripes on the back, and one on each 
side of the body. The glands were usually darker than the surrounding skin, 
and in some examples scattered glands bore brick red caps, which with darker 
rings appeared as ocelli. Some specimens were lighter than others, and an oc- 
casional one had a strong infusion of pale reddish brown" ( J. O. Snyder, Nev., 
1920, pp. 83-84). 

"The colour in life was grayish green above and more or less mottled; a 
longitudinal whitish stripe down each side of the back from behind orbits to 
a little behind knees, and one along each side from behind ear; a short median 
whitish stripe on posterior part of back (over urostyle) ; a darkish blotch over 
orbit passing backward and inward, and a darkish line from eye to nostril; 
tubercles on back and sides tinged with orange; throat dusky, belly white" 
(E. B. S. Logier, B.C., 1932, p. 319). 

Structure: "No interorbital boss present. In some specimens of intermonta- 
nus there is a glandular interorbital elevation which resembles the true boss 
found in bombijrons. This may be removed and the true nature of the skull 
revealed. . . . Head width wider 20.9-22.5 mm. Body rugose or with many 
individual prominences or warts. Color mottled whitish and black above; ven- 
ter whitish; in preservative the back becomes blackish with some white areas. 
At times the back is streaked with whitish lines. Venter white. No fronto- 
parietal fontanelle; interorbital space with prominent fronto-parictal bones 
forming ridges; ... in some specimens the interorbital space is filled with a 
glandular prominence resembling the bombifrons species; head width 22.5 
mm., whole foot 31.2, confined in the main to the Great Basin area" (V. M. 
Tanner, Gen., 1939, pp. 11-12). 

Voice: "Their appearance was at once announced by a loud chorus which 
differed markedly from that of Hyla or Rana, being in a lower key, somewhat 
guttural, and a little rasping. It was entirely different from that of Bufo" (J. O. 
Snyder, Nev., 1920, p. 83). 

"I would describe the call as a soft, though very penetrating \wa\, low- 
pitched with something of the quality of the vibrating of a heavy rubber 
band" (W. Wood, Utah, 1935, p. 101). 

"A very secretive, wholly nocturnal, but really common species. Its call, a 
loud crah-crah-rah, repeated at short intervals, was first heard at Bcllevue late 
in April and again, following a cold spell, on May 5. Thereafter it increased 



122 HANDBOOK OF FROGS AND TOADS 

in volume nightly until June i, after which it decreased, stopping about June 
15. Rain puddles, overflows from irrigation ditches, in fact every pool of stand- 
ing water served as breeding places. Drying up of the pools rarely permitted 
a development beyond the egg or early larval stages" (G. P. Englehardt, Utah, 
1918, pp. 77-78). 

Breeding: Records indicate breeding from April (Hardy), May (Snyder, 
Carl), June (Snyder, Tanner, Wood), July. Transformations are from late 
May through June and July to August or September. 

"This, the commonest Carbon County amphibian was heard croaking on 
April 9, 1937; eggs were collected April 22. Numerous; breeding near river at 
Price, April 27. Also found breeding in puddles of rain water, in stagnant 
pools, in water, in alkali washes, in city reservoirs and swimming pools, and in 
sewage near Price River. Eggs deposited in the laboratory hatched after four 
days. Eggs collected in water containing algae were green due to inclusion of 
algae in the outer jelly while eggs deposited in muddy water included soil" 
(R. Hardy, Utah, 1938, p. 99). 

"On the evening of June 2, 1911, 1 happened upon a small pond separated 
from the water of Pyramid Lake by a narrow bar. The pond was but a few feet 
in width, and perhaps a hundred feet long. The water was clear and slightly 
alkaline like that of the lake. In it were hundreds of spadefoots depositing 
their eggs in masses one layer deep on the upper surface of small rocks. The 
eggs were not piled up after the manner of frogs, nor were they in strings like 
those of toads. One mass presented fresh eggs and likewise others in which 
development was marked, plainly indicating that the mass was made of at 
least two contributions" (J. O. Snyder, Nev., 1920, p. 84). 

Tanner records and describes a 5i-mm. tadpole, coppery color in water, 
bluish black in preservative. He gives labial teeth rows as %. Snyder secured 
a 6o-mm. tadpole with teeth %, % and transformation sizes of 20-24% mm. 

"From accounts of spadefoots that I have read I judged that the toads come 
out of their burrows only after a rain, and that a certain amount of moisture 
was necessary to bring them to the surface. However, the fact that so many 
hundreds of individuals came out of the ground in Smoky Valley in 1932, 
when no rain had fallen for a long time, indicates that other circumstances arc 
able to initiate activity in this species." "In 1933, with Compton I worked in 
this same vicinity from mid-May to mid-June. The pasture was fairly dry this 
time, but a large proportion of the 1932 brood of spadefoots had survived, and 
we saw them almost daily. We were surprised to see them day after day forag- 
ing on the surface of the ground in daylight hours" (J. M. Lmsdale, Nev., 
1938, pp. 21, 23). 

"On July 4, many small, recently transformed specimens were collected in 
mud cracks in the bed of a dricd-up irrigation ditch near Maggie Creek. It 
was found that the easiest method of collecting them was to stamp on the 
ground. The vibration disturbed them and they would thrust their heads out 



SPADEFOOTS: SCAPHIOPODIDAE 



123 



of the cracks. If the jar continued, they came out and hopped about on the 
ground where they were conspicuous and easily captured, but when the 
stamping ceased they soon disappeared. A few had completely transformed, 
but the majority had tails varying in size from mere rudiments to the length 
of the head and body. Later many half grown specimens were observed com- 
ing up out of the ground behind the mowers in a hay field near Annie Creek" 
(A. G. Ruthven and H. T. Gaige, Nev., 1915, p. 16). 

Journal notes: May 20, 1942, Beaver Dam Lodge, Ariz., at bridge. We heard 
in several directions a few Scaphtopus hammondii subspecies. They were 
mainly concentrated in the big pool at north end of bridge. Two were too far 
out to collect only one did we get. Its call is a harsh far-carrying call very dif- 
ferent from the trill of B. woodhousn. 

June 5. Stopped at Jacob's Lake, Ariz. The assistant forester, Harlen John- 
son, said about May i they heard an awful squawking and he wondered what 
it was. I asked him if they had spadefoots. He said "Yes." He sent droppings 
of coyotes to the Colorado laboratory and they said there were spadefoot re- 
mains in them. I went down to the lake and found two sizes of Hammond's 
spaclcfoot tadpoles. 

Authorities 9 corner: 
E. B. S. Logier, B.C., 1932, p. 319. W. F. Wood, Utah, 1935, pp. 101-102. 

Spadefoot, Holbrook's Spadefoot, Hermit Spadefoot Toad, Spadefoot Toad, 
Hermit Toad, Solitary Toad, Solitary Spadefoot, Hermit Spadefoot, 

Storm Toad 

Scaphioptts holbroofyi holbrookji (Harlan). Plate XXI; Map 7. 

Range: From West Virginia (Green and Richmond, 1942) and Ohio, down 
the Ohio River to southeastern Missouri and northeastern Arkansas, across 
extreme southeastern Texas, and along the Gulf and Atlantic coasts to Massa- 
chusetts. 

Habitat: Shallow burrows in the ground; nocturnal in habit. They breed 
only after heavy rams, and then usually in temporary rain pools. The follow- 
ing note relates several items about their general habits. Dave Mi/zell of the 
Folkston, Ga., region, told of "a frog called 'storm frog' because it appears 
when there are storms or floods. It hollers wanl(! It folds up so that you can't 
see its legs. It is larger than the 'hop toad.' One was dug up in a potato field." 
"Very common at breeding time; seen on warm nights at the mouth of bur- 
rows, and hopping about in the woods" (O. C. Van Hyning, Fla., 1933, p. 3). 

No other form in this country has had more printed about it than this elu- 
sive form. Our account, however, will be one of our shortest summaries. No 
one has nightly visited these creatures in their nonbreeding holes more than 
Neil Richmond. 

Size: Adults, 2-2% inches. Males, 54-72 mm. Females, 50-71 mm. 



I2 4 



HANDBOOK OF FROGS AND TOADS 



General appearance: Like the common toad, it is short and broad in body, 
and with small round posttympanal glands (parotoids) . The skin is relatively 
smooth but bears scattered warts; is usually brown in color, frequently with 
two more or less evident light dorsal stripes. The arms and legs are short and 
thick; the feet, broad. The inner tubercle of the sole is a large dark horny 
process with which the toad digs its burrows. The eyes are large and protu- 
berant with vertical pupils indicating nocturnal habits. The throat and breast 
are white; the lower belly is grayish. 




Map 7 

Color: Milliard, Fla., Aug. 18, 1922. Male. Stripe from eye back to vent 
lemon yellow or greenish yellow; other light colors of the back are same as 
stripe from eye to vent, so also on the sides. The snout is mummy brown, 
Prout's brown, sometimes blackish brown, or bone brown. This is also the 
remaining color of the back, which is sometimes virtually black. Limbs on 
dorsum, much the same color or slightly washed out greenish yellow of the 
light back colors; under parts of hind limbs and lower belly light grayish 
vinaceous. Rest of the under parts are white, especially the white glistening 
throat, which shows so conspicuously in the "croaking bubble." First three 



SPADEFOOTS: SCAPHIOPODIDAE 



125 



fingers with black excrescences. 
Spade of hind foot dark-edged, as is 
also the tip of first toe. Web of foot 
much darker than the grayish vi- 
naceous web of female. Iris light 
greenish yellow with black on the 
outer rim. Pupil vertical. 

Female. Sometimes uniform 
warm sepia or bone brown above. 
Throat and breast white. Underside 
of fore limbs and hind limbs and 
lower belly light grayish vinaceous. 
Sometimes the stripe on females 
may be sulphur yellow instead of 
the intense greenish yellow of the 
male. Almost always each male of 
a pair had a predominance of yel- 
lowish hues and the females in- 
clined to the brownish hues. 

Structure: Tympanum much 
smaller than the eye; large, wide 
hind feet; skin on crown of head, 
thin; parotoid gland present; male 
with a subgular vocal sac; males 
with fingers broader than females. 

Voice: Hoarse, coarse, monosyl- 
lable, waftl{, wanl{, like the calling 
of young crows. Aug. 16, 1922, we 
heard the congress at a half-mile dis- 
tance. At this distance, to one it 
sounded like the calling or snarling 
or complaining note of a cross baby; 
to another member of the party, like 
young crows; or at other times like 
young herons in a herony. One 
member characterized it as naarh 
naarh, complaining, nasal, not shrill 
or high-pitched. Near by it sounds 
somewhat like where, where, where, 
where. 

When we first approached, the 

males called on all sides in full daylight, almost at our very feet. The inflated 
throat by day is a beautiful glistening white golf ball. The male before he calls 




Plate XXL Scaphtopus holbroo^ti hoi- 
brookii. 1,4. Males (X%)- 2,3. Females 
(X%) 5- F -gg mass (X 1 /^- 6- Forearm of 
male (X%). 



126 HANDBOOK OF FROGS AND TOADS 

lies on the water's surface with hind legs partially submerged. When he croaks 
he dips the hind end of the body and the head is reared to a 45-75 angle 
with the water's surface. When at the height of the performance, or slightly 
before, he closes his eyes. Then the throat deflates and the body inflates. He 
croaks about once in every two seconds. 

A big congress may make as much noise as a steam calliope, though of a 
different nature. Normally the mated pairs are quiet. It is the calls of free 
males which are searching a mate or annoying the already mated pairs which 
make up the chorus. 

"The peculiar, harsh croaking of this singular toad must be heard to be 
appreciated, and can then never be confounded with that of any other species. 
The only sound we can liken it to is that of a heavily loaded, creaking wagon 
rolling over hard and uneven ground" (F. W. Putnam, Mass., 1867, p. 
109). 

"We have found only one reference to a spadefoot singing while under- 
ground (A. H. Wright, 1932). We found this to be one of the most striking 
peculiarities of the animal. In the early part of the evening, between nine and 
ten o'clock, no toads were visible in the ponds, but from all around the margin 
came their much muffled calls. It was a most peculiar sensation to be in the 
midst of the chorus but to have only a barren expanse of sticky clay visible un- 
der the flashlight. Excavating with considerable difficulty we found the song- 
sters to be several inches below the surface in apparently quite solidly packed 
earth. No sign of entrance to the burrows was visible. As the evening wore 
on one after another pushed out to the surface and entered the pond to float 
and sing with greater vigor. The chorus reached its height shortly after mid- 
night" (E. G. Driver, Mass., 1936, pp. 67-68). 

Breeding: On this topic as on voice there are countless observations. The 
spadefoot breeds from March to September at periods of heavy rainfall. The 
eggs are in irregular bands along grass blades or plant stems, the band 1-2 
inches (25-50 mm.) wide and 1-12 inches (25-300 mm.) long, the egg 1 /ic- 
Ki> inch (1.4-2 mm.), the envelope %o-/u inch (4-5.6 mm.). They hatch in 
i 1 /i>-2 days. The bronzy tadpole is small, i% inches (28 m.) broad, but not 
deep, its tail short and rounded. After 14-60 days, the tadpoles transform from 
July to September at %-% inch (8.5-12 mm.). 

Journal notes: Aug. 16, 1922. We started from Callahan in a hard rain, a 
little before noon. On a detour two miles south of Hilliard, Fla., we stopped 
for cars going across a swollen creek. Francis went to look for birds and heard 
spadefoots calling. We drove through the woods and back on the Dixie High- 
way and pitched camp, i mile south of Hilliard on an oak ridge. An old road 
filled with water made a shallow pond, and here we saw the males croaking, 
their white throats looking like shiny white golf balls. Just beyond was a 
shallow surface pool made by the heavy rains. The ground was covered with 
herbs: a little Xyris, a few sedges, Rhexia, a small umbelliferous plant with 



SPADEFOOTS: SCAPHIOPODIDAE 



127 



violet-shaped leaf, wire grass, and Hypericum. The spadefoots were calling 
here, in another similar pool, and in a third deeper pool as well. At a distance 
the chorus sounded like young crows trying to call. The pond was filled with 
mated pairs. 

A few eggs had been laid. The eggs were laid in more or less irregular band 
form along the grass blades or plant stems. In the third pond where the water 
was deep, the bands were long. The pair might be floating on the surface. 
When ready to lay, they went to the bottom of the pond, often the male with 
his eyes closed and the female with hers partly closed. They moved along 
slowly on the bottom or rested a minute. When she found a stem to suit her, 
she seized it with her front feet and pushed with her hind feet. The male clung 
close to her back, his chin tight against her back. (One we photographed had 
an abrased chin as if from pressure.) He held his knees against her knees, or 
sometimes his feet, which are conspicuously broad, were pressed against her 
feet. She walked or climbed up the grass blade or along it, if it fell to hori- 
zontal position, and pressed her vent against the blade as she laid the eggs. He 
humped his back to press his vent close to hers while she was laying. As they 
reached the top of the blade, they sometimes moved immediately to a nearby 
one, or rested a short period. When first laid, the eggs had an irregular band 
appearance as they were strung along the blade, sometimes being much 
thicker if more eggs had been emitted at such periods. When first laid, they 
had a brownish appearance with conspicuous creamy-white vegetative pole. 
As the jelly swelled and the eggs all turned right side up, they looked very 
black. 

By the next morning some eggs were almost ready to hatch (these must 
have been the ones we found when we first found the pond). The other clus- 
ters were swollen into loose, irregular, elongate bunches attached to the stems 
which tipped so that many times the bunches lay lengthwise on the water. 
There seemed to be a tendency for the bunches of eggs to be more or less clus- 
tered in areas. We noticed many pairs close together that first afternoon, and 
unattached males trying to get to a female, thus making tangled masses of 
toads. There was a strong chorus that night and by the next morning the pond 
was all churned up and muddy. Many, many eggs were there, but no toads. 
The story was told for the season. 

Authorities' corner: 

A, Nichols, Mass., 1852, pp. 113-115. F. Overton, N.Y., 1915, p. 17. 
F. W. Putnam, Mass., 1865, p. 229. N. D. Richmond, Va., 1947, pp. 53-67. 

Key West Spadefoot 

Scaphiopus holbroo1(ii albits (Carman). Plate XXII; Map 7. 

Range: Florida Keys and possibly the extreme southern part of Florida. 
Habitat: The query that naturally arises is: Do the other species of frogs 



128 



HANDBOOK OF FROGS AND TOADS 



such as the green tree frog, southern tree toads, toads, or frogs have a similar 
tendency toward albinism in the Keys? Is it due to salt or a light beach habi- 
tat? Or, was this lot of spadefoots an isolated albinistic collection that might 
possibly occur on the mainland in other portions of the spadefoot's range? 
One would expect the subterranean spadefoot to tend more toward albinism 
than almost any other species of the country. 




Plate XXII. Scaphiopus holbrool{ii albus (XO- i> 2 >3- Males, trom UbJNM. 
4. Male, from Gainesville, Fla. 5. Female, from Gainesville, Fla. 

Size: Adults 2%-2% inches. The three specimens in the National Museum 
are males measuring 54, 56, 56 mm. The University of Michigan Museum has 
a female 56 mm. 

General appearance: This spadefoot is like Holbrook's but with an exces- 
sive amount of white in the pattern. 



SPADEFOOTS: SCAPHIOPODIDAE 



129 



Color: "Average size less than that of preceding (S. h. holbroo\ii). Brown 
of the back lacks the red or chocolate tinge. Readily distinguished by the great 
amount of white on back, flanks, and upper surface of limbs. The white forms 
spots or vermiculations which coalesce into bands of irregular shape and ex- 
tent" (S. Carman, Gen. Check. L., 1884, p. 45). A doubtful subspecies. 

Structure: Apparently the interorbital distance is narrower in S. h. albus 
than in S. h. holbroofyi, being in body length 9.3-10 in S. h. albus and 6.77-8.5 
in S. h. holbroo\ii. Two pectoral glands are present. 

Scaphiopus h. albus holbroo\ii 

Male Female Male Female 

Body length 56mm. 56mm. 56 mm. 56mm. 

Interorbital space 6.0 mm. 5.5mm. 8.0 mm. 8.0 mm. 

Voice: Their cries sound like "ow, ow," and "miow," but the latter much 
deeper in tone than the well-known cat cry. The noise made by a dozen males 
is deafening when one is near, though the call lacks carrying power (R. F. 
Deckert, Fla., 1921, p. 22). 

"The habits of the two seem to me to be identical, and I can detect no dif- 
ferences in the voices" (A. F. Carr, Jr., Fla., 1940^ p. 54). 

Breeding: We have seen live specimens from Gainesville which seemed 
almost as light as the preserved specimens of this subspecies. Pending the de- 
termination of the status otS.h. albus and whether it extends to the southern 
tip of Florida, we wish to point out that Richard F. Deckert does not describe 
the spadefoots of Miami as S. h. albus. He calls them Scaphiopus holbrootyi. 

"Scaphiopus holbroofyi (Harlan). On April 23 (1920) a male specimen was 
found by the writer, about six inches down, in sandy marl at Brickell Ave. and 
Broadway, Miami. During a prolonged thunderstorm many of the spadefoot 
toads were encountered by the writer on the streets south of Miami River, on 
the afternoon of May 16, and during the night were found breeding at igth 
Street and Ave. H., also at 22nd Street and Miami Ave., and great numbers 
were reported from the low grounds near the 'Alligator farm/ Miami" (R. F. 
Deckert, Fla., 1921, p. 22). 

Journal notes: March 18, 1934. Looking for Florida spadefoots, we spent one 
night at Caribee Colony on Matecumbe Key, and went out in the evening 
with our flashlights to "shine their eyes." A few were out, but more were in 
their burrows with the tops now shaken open. 

Authorities' corner: 
C. E. Burt, Gen., 1938, p. 335. 
V. M. Tanner, Gen,, 1939, p. 3. 
A. F. Carr, Jr., Fla., i94ob, pp. 31, 53-54. 



130 



HANDBOOK OF FROGS AND TOADS 



Hurter's Spadefoot, Hurter's Solitary Spadefoot 

Scaphiopus holbrootyi hurtcrii (Strecker). Plate XXIII; Map 7. 

Range: Eastern half of Texas. Records exist from Houston and Edna to 
Cameron County. Smith extends it to western Arkansas. 

Habitat: Like other spadefoots, these frogs come out of their burrows to 
breed in temporary pools. 

Size: Medium. Type 67 mm., from Waco. Refugio specimen 63 mm. The 
range of size of ten breeding adults from Lytle, Texas (collected by A. J. Kirn, 
June 28, 1931) is 66-78 mm. Adults, r%-3% inches. Males, 43-73 mm. Fe- 
males, 44-82 mm. 




Plate XXIIL Scaphtopus holbwolyi hurtem (X%)- i>23>4>6. From Albert 
Kirn, Somerset, Tex. 5. From USNM to show pectoral glands. 

General appearance: "Head short, length about equal to width. (In hoi- 
brooch the head at angle of mouth is much wider than long.) Snout heavy 
and blunt, not extending beyond the mouth. Parotoids nearly round, higher 
and even more conspicuous than in the eastern species. Tympanum distinct 
but rather smaller than in holbroofyi. (In type hardly more than half the 
diameter of the parotoid.) Crown distinctly rugose. No black granules in 
space between and in front of the eyes. Upper surfaces with small, closely set 
tubercles, very uniform in size and distribution. Many tubercles on sides, but- 
tocks, and posterior portion of the abdomen. 

"Color above, pale greenish, with a pale yellowish line from each orbit; 
these converge again in the coccyx. Upper surface of head and area between 
the light lines, dark plumbeous, parotoids olive. Sides of head and under sur- 
faces yellowish-white" (J. K. Strecker, Jr., Tex., 1910, pp. 116-117). 

Color: Somerset, Tex., from Al Kirn, June 20, 1945. Males. The dorsal back- 



SPADEFOOTS: SCAPHIOPODIDAE 131 

ground color is Saccardo's olive, medal bronze to raw umber. A light stripe on 
either side crosses the rear of the eyelid, outlines the corner of the head boss, 
the pair becoming parallel for % inch and about % inch apart, next bowing 
outward and then approaching each other toward the groin. These stripes arc 
dull green-yellow or bright chalcedony yellow to buffy citrine. From the vent 
forward is a short middorsal stripe of the same, which also occurs on outer edge 
of foot and tarsus and on a few tubercles on the outer edge of tibia and along 
the sides of the body. There is a patch of the same color above the arm inser- 
tions. The tympanic area is buffy citrine to dull green-yellow. The top of head 
and boss are smooth and unspotted mummy brown. The iris is light cendre 
green to light ochraceous-buff with a horizontal bar of black, which is seldom 
complete. The under parts are cream to white, most intense on chin and pec- 
toral region. The rear portions are vinaceous-buff, which color sometimes suf- 
fuses the whole under parts. The upper % of first finger and inner and upper 
half of second bear black excrescences. 

Females. Females have a tendency to be less green. The stripes are more 
buffy, in one almost white. The lateral tubercles may be almost white and 
arranged in 5-7 series. The excrescences on fingers arc lacking. 

Structure: The boss on head is large, conspicuous, and much raised at the 
rear. The axillary breast glands are present. 

"Many pustules on upper surface of tibia. Glands on thorax present, con- 
spicuous. Enlargements resembling glands on inferior surface of femur (pres- 
ent in both specimens). Spade-like process of foot narrowly margined with 
black. Palmar tubercles rather small. Fingers slender. Tibia about equal to 
that of S. holbroo^ii but femur and foot much shorter" (J. K. Strecker, Jr., 
Tex., 19103, pp. 116-117). 

Distinguished from S. h. holbroofyi by "its more compact form, narrow 
head, blunt muzzle, unusually high parotoids, smaller palmar tubercles, and 
shorter hind limbs. The sides, buttocks, tibia, and posterior portion of the 
abdomen are covered with tubercles instead of being almost perfectly smooth. 
The tubercles on the upper surfaces are more uniform in size" (same, p. 116). 

Mr. Kirn's material (June 28-29, 1931), when compared with male S. h. hol- 
brootyt of the same size, has smaller measurements. The head to angle of 
mouth, the width of head, the tympanum (equal in one), the snout (greater 
in one) were less than in S. h. holbrootyi. The hind limbs were equal in the 
two species. 

Mr. Kirn's specimens include 66-, 68-, 68-mm. males. We studied also Strcck- 
er's material and that of USNM and Baylor University. Our custom of meas- 
uring males and females of every species of frog in the United States at 20, 28, 
36, 44, 56, 66, 68, and 82 mm. makes the measurements more readily compara- 
ble than random ones of unscxed specimens. The following table is part of 
the 25 measurements we made for each specimen: 



132 HANDBOOK OF FROGS AND TOADS 





5. h. 


S. h. 


S. h. 


S. h. 




hurteni 


holbroo^ii 


hurterii 


holbrool(ii 




44 mm. 


44 mm. 


56 mm. 


56 mm. 


Head to angle of mouth 


H 


*5 


15 


18 


Width of head 


*7 


19 


21 


23 


Snout 


8 


8.5 


9 


ro 


Tympanum 


3 


4.0 


3-5 


5-5 


Hind limb 


43 


45 


53 


49 


Tibia 


14 


'5-5 


18 


17-5 


Foot with tarsus 


22 


24 


28 


28 


Foot without tarsus 


16 


16.5 


20.5 


i9 



Voice: "The call is a single note, which, while guttural, has a peculiar soft 
quality not unpleasing to the human ear, quite different from that of Scaplno- 
pus bombifrons, S. cotichii, or 5. hamrnondii, and different also from descrip- 
tions of the cry of S. h. holbroolyi, which is commonly stated to be very harsh 
and loud (cf. Ball, 1936). Nevertheless the voice of hurteni has a quality char- 
acteristic of spadefoot breeding calls; I recognized it at once when first heard 
as a spadefoot call. The cry is not exceptionally loud, although a large chorus 
can be heard for at least one-half mile. (I have heard S. bombtfrons for more 
than 2 miles on a still prairie night.) The calls of males tend to stimulate other 
males to call and to attract both females and other males to a given pool. 

"Each call is explosively given, and the vocal sac becomes fully distended 
and then deflated each time. Intervals between calls vary, both with the num- 
bers of individuals present and with numbers calling at any one time. The 
usual interval m a large, excited congress is from one to two and one-half sec- 
onds. There is no tendency for the calls of males to be synchronous" (A. N. 
Bragg, Okla., 1944, pp. 231-232). 

Breeding: January to December (Kirn). 

"All masses of eggs of S. httrtern seen were produced near the shore, at and 
near the water surface, and strung out over vegetation. When in place, they 
superficially resemble those of Bttfo rather than those of other spadcfoots. Each 
egg is black at the animal pole, shading to very light grey or white at the vege- 
tal. Each is inclosed in a ball of jelly considerably larger than itself, i.e., in a 
single thick gelatinous envelope. Individual eggs vary little in size, each being 
about 2.3 mm. in diameter. The gelatinous capsules also are quite uniform at 
about 6.7 mm. The gelatinous coats are elastic and quite sSticky so that they 
easily pick up particles from the surrounding water. This tends to conceal 
them after a few hours. The little gelatinous balls tend to stick together in 
various patterns. These patterns are quite irregular, but most often the eggs 
attach tandem-fashion so that an irregular string is produced with some simi- 
larity to the eggs of toads. There is no continuous, gelatinous, encasing tube 
as in Bufo, and one egg within its covering can easily be separated from others 



SPADEFOOTS: SCAPHIOPODIDAE 



133 



to which it is joined. Such strings may be very short or several inches long. 
Individual eggs of one such string commonly attach to others in adjacent 
strings, forming a network. Several such lattices are sometimes similarly 
joined to form a three-dimensional pattern. A winding string of eggs, loosely 
attached to other such strings wherever contact happens to be made between 
them, is most commonly found. 

"The number of eggs in a complete clutch is unknown, but the complement 
must consist of several hundred at least" (A. N. Bragg, Okla., 1944, pp. 232- 



Journal notes: On Feb. 9, 1932, Mr. A. J. Kirn of Somerset, Tex., sent us ten 
preserved Scaphiopus which Mr. Strecker pronounced S. h. hurterii. On Feb. 
15, 1932, Mr. Kirn wrote, "They [the spadefoots] are not calling as yet. I 
found one of them last year, January 27 in the road, during a rainy spell. 
I will send you some of the Scaphiopus when they come out to lay. Do not 
know when this will he." His field notes for these ten spadefoots are some- 
what as follows: "Lyile June 29 (Monday) 1931. Rain Friday 8 P.M. through 
most of Saturday and again yesterday, early morning and late evening, again 
early this morning and this afternoon, about three inches altogether. Weather 
warm. Scaphiopus hurterii heard last night at all pools at wells (old slush 
pits). Collected a dozen at No. 10 and No. 13 (4 pairs in copulation; in all of 
these the smaller and lighter one was male). Collected between 10 and n P.M., 
June 28. This afternoon, I found eggs already hatched at Nos. 10 and 13 wells. 
No spadefoots heard before 8:30 P.M. yesterday. Eggs evidently laid last night 
after midnight. Were hatched at 7 P.M. today and how long before I do not 
know. The spadefoots collected last night varied from gray and yellowish 
green to dark, all with two dorsal stripes (widening on middle posterior 
back). Parotoid glands distinct in all. Had in can all of today. Many eggs laid 
m the can. Not any spadefoots heard or found tonight, June 29." 

We had a group of spadefoots in a laundry tub, in which we had $ inches 
of dirt. We fed them mealworms, from a glass dish in the center of the tub. 
By clay no toads were visible, nor was there any evidence of their burrows, 
but about 9:30 in the evening they began to shake their heads and loose spots 
appeared here and there, and slowly the toads came out, and quickly snapped 
up the proffered worms. 

We have no personal experience with this form in the field. Our latest lot 
(1945) of live material came from A. J. Kirn, who knows it best. Some of his 
notes are: 

"2/12. Much rain last 2 nights. Spadefoot toads calling loudly tonight 7:30 
on. From water pools. This first heard this spring and winter. Weather mild. 
2/12. Very heavy rain last night and nearly as much night before that. Many 
spadcfoot toads calling from all pools. Temperature 57 at H P.M. 2/13. Egg 
clusters of spadefoot toads this A.M. in a pool. Also numbers of tadpoles, vari- 
ous size from small (% inch) to large, hind legs showing beneath skin. Evi- 



I 3 4 HANDBOOK OF FROGS AND TOADS 

dently eggs laid some time ago. (January?) See 2/21. 2/17. A few spadefoots 
in roads tonight. 2/21. Tadpoles, large, of 2/13 have hind legs today. Egg 
cluster pool of 2/13 has numbers of tadpoles about 6 mm. 2/27. Spadefoot tad- 
poles of Feb. 12-13 still not as large as the largest seen Feb. 13. Therefore those 
were from eggs laid in January, possibly about January 21. This is my first and 
only record of eggs laid during that month, though I always contended that 
they would lay eggs any month of the year if temperature and conditions were 
right. Weather colder Feb. 26 and 27. March i. Milder and foggy. 

"7/5. On June 19, 1 sent you a box with 25 spadcfoot frogs, hope that they 
reached you alive, and that they were enough to suit your needs. We had a 
four-inch rain the night before and they called until 12:15 noon, then sun came 
out, they quit until after sundown, when some came out. I went to nearby 
ponds (tanks) found them scarce, and wild. However I managed to pick up 
two dozen, then got one in the house-yard. Can get more later on when it rains 
hard enough, if you need them." 

Authorities' corner: 
II. M. Smith, Tex., 1937, pp. 104-108. 
V. M. Tanner, Gen., 1939, p. 9. 
A. N. Bragg, Okla., 1942!^, p. 506. 



FAMILY BUFONIDAE 

Genus BUFO Laurent! 
Maps 8-14 

Colorado River Toad, Giant Toad, Girard's Toad, Colorado Toad 

Bufo alvarius Girard. Plate XXIV; Map 8. 

Range: Imperial Valley, Calif., up the Colorado River almost to Nevada's 
tip, up the Gila River almost to New Mexico's border, and south into Sonora, 
Mex. 

Habitat: Lower Sonoran life zone. Semiaquatic. In the general locality of 
large permanent streams or the irrigated portions of our southwestern desert 
regions. 

Pima Co., Ariz. "Bufo alvarius is much commoner near the Steam Pump 
than other species of the genus. With a few exceptions it is found only in the 
wet places around the cattle watering troughs of the ranches in the mesquite 
association. The only other place where we found it was in a temporary road- 
side pond formed by a heavy rain where Scaphiopus couchii and S. ham- 
mondti were breeding. These, however, had very probably come from the 
watering troughs of the ranch only a few hundred yards distant. While Bufo 
alvarius was found in the pond with the breeding Scaphiopus, none of the 
former were breeding. One specimen was found at dusk at the mouth of a 
canyon on the dry sandy bed, and a few were found at night by the roadside. 
Practically all were found after dark, but we discovered that their daytime 
hiding place was in hollows under the watering troughs" (A. I. Ortenburgcr 
and R. D. Ortcnburger, Ariz., 1927, p. 102). 

Size: Adults, 3M-7 inches. Males, 80-156 mm. Females, 87-178 mm. 

General appearance: This is a very large grayish or brownish-green toad 
with smooth, leathery skin and a few scattered small, rounded warts. The un- 
der parts are light. The head is broad and flat, marked by low, broad, crescent- 
shaped crests curving around the rear of the eye. These are like fleshy folds. 
The canthus rostralis is marked by a ridge which turns down in front of the 
eye as a preorbital. One to four white warts are present back of the angle of 
the mouth. The parotoids are large, subreniform in shape, spreading down- 
ward at the shoulder. There is a large glandular wart on the femur and a long 
one or several shorter ones on the tibia. These glands are the conspicuous mark 



136 



HANDBOOK OF FROGS AND TOADS 



of this toad and appear early, being present in a r%-inch (44-mm.) individual. 
Color: Female. Bard, Imperial Co., Calif., from L. M. Klauber, May 22, 
1930. Color deep olive, dark olive or citrine-drab, or dark greenish olive on 
back and buflfy olive on either side of back. Warts on back not conspicuous, 
buckthorn brown or Dresden brown. Some of same color on each upper eye- 
lid. This brown scanty on femur, absent on forelegs, tibia, and hind foot. Fore 
limbs deep grayish olive. Arm insertion olive-gray and some spots of same 
color each side of pectoral region and edge of throat. Throat same color as 
venter. Venter white. Back of angle of mouth are two pale ochraccous-buff or 
shell pink glands. Tympanum like dorsal color. Iris cream-buff below and 




MapS 

above, with some tawny or russet streaks. Dark longitudinal bar through eye, 
brownish olive ahead of and behind eye. 

A young specimen (USNM no. 21802, 44 mm.) has a spotted back and at 
first appearance looks like a B. punctatits. The parotoicls are oblong, spreading 
apart, the glands on femur arc close to tibia, and those on tibia barely show. 
Tympanum very distinct, slightly elliptic. Two white warts back of angle of 
mouth present. 

Structure: Two metatarsal tubercles; two large palmar tubercles; first finger 
about equal to second; first finger of female may look very long and slender, 
that of male much heavier at base; palms and soles tuberculate; interorbital 
much wider than internasal space; a membranous fold at the inner edge of 
the tarsus; horny excrescences on fingers of male may be very prominent, 
starting from back of wrist and extending all along inner side of first finger 



TOADS: BUFONIDAE 



137 



and covering upper surface as well; second finger has upper surface with ex- 
crescence as well as triangular patch from tip backward; slight on third; tym- 
panum may be obliquely vertical and elliptical or almost round and very little 
oblique. 

Voice: "When held in the hand, this toad jerks spasmodically, and vibrates 
the whole body, as if about to explode with wrath. The only sound, however, 
produced in protest is a gentle chirping note, less loud and emphatic than that 
of the American toad" (M. C. Dickerson, Gen., 1906, p. 108). 

"I assure you there was no lack of noise that day nor night, the croaking 
being incessant" (A. G. Ruthven, quoting J. J. Thornber, Ariz., 1907, p. 506). 

"On the night of July T3th, a chorus of giant toads was heard from a ditch 
near the Sells-Roblcs road southwest of Tucson" (C. F. Kauffeld, Ariz., 1943, 

P-343)- 
"Of batrachians, a toad (Bufo) and a frog (Runa virescens brachycephala 

Cope) were found at Warsaw Mills; and at Buenos Ayres, at the beginning 
of the summer rains, Lieutenant Gaillard observed great numbers of a very 
large froglike toad, named Bufo alvarhts by Girarcl. Nothing was seen or 
heard of them until the advent of the early summer rains, which formed a 
large shallow lake near Buenos Ayres and about 10 kilometers (6 miles) north 
of the Boundary Line. These large toads then filled the air with their loud 
cries, which increased until a deafening roar was produced. Numbers of them 
were seen hopping about, but their rarity was not suspected by Lieutenant 
Gaillard, on which account none were collected" (E. A. Mearns, Ariz., 1907, 

P-"3) 
Breeding: "Deep brown above and tan below, the eggs arc encased in a 

single long tube of jelly. The envelope is somewhat loose, but quite distinct 
in outline; the gelatinous material is, for the most part, clear, transparent, and 
not very adhesive. There arc no partitions between the individual eggs, whose 
arrangement varies from a perfectly linear scries of near spheres to a zigzag 
pattern of broadly wedge-shaped eggs. The number of eggs per inch averages 
eighteen (range 12 to 28). Vitcllinc membranes arc close to the vitclli and not 
visible with the unaided eye; at times they are difficult to see even under mag- 
nification. Measurements on the eggs are as follows: vitellus 1.4 mm. (range 
1.14 to 1.70 mm.) ; vitellme capsule 1.6 mm. (range 1.25 to i .70 mm.) ; envelope 
2.2 mm. (range 2.12 to 2.25 mm.). Affinities: Except for the deeper color of the 
vitelli, the lesser degree of convolution and lack of slight scalloping of the 
gelatinous tube, eggs of B. alvartus are similar to those of B. compactihs. Meas- 
urements of both average approximately the same m all respects. The total 
mass of eggs is larger in B. alvariits and the crowding within the tube is more 
intense (12 to 28 per inch for alvarius to 14 to 20 for compactilis), this differ- 
ence, however, being barely noticeable. Superficially B. alvarius eggs also look 
like those of B. terrestris. But here the lighter color of the eggs and greater 



138 HANDBOOK OF FROGS AND TOADS 

degree of convolution of the tube plus the presence of an inner envelope serve 
to separate eggs of these species immediately." (R. Livezey and A. H. Wright, 
Gen., 1947, pp. 193, 194,214). 

The eggs we secured July 2-3, 1934, were * n l n & ropelikc strings (400 
inches) of 7500-8000 eggs. 

"Two of the females collected at Alamos between August 27 and Septem- 
ber 2 contain mature eggs in the oviduct" (C, M. Bogert and J. A. Oliver, 
Gen., 1945, p. 339). 

Journal notes: July 30, 1917, just southeast of Tempe, Anz. Made only 49 
miles. In a water hole near a culvert Ralph Wheeler and I caught six immense 
toads (Bufo alvanus). All males. Probably tardy ones. Tadpoles (Bufo) in the 
hole probably of this species. 

June 25, 1934, Florida Experiment Station, Ariz. Mr. R. R. Humphrey said 
that at 10 P.M. on the road from Nogales north he saw several Bufo alvanus 
after rain. Mr. Gorsuch said they often gather at the Continental pond, that he 
once found several pairs of B. alvanus which he brought up to the station. 
They laid long strings of black eggs. Had he known they were not described 
he would have saved them. 

June 29. At Continental we walked around an irrigation pond. Under a tin 
cover found a large male B. alvanus. How he did protestingly chuckle as we 
carried him off. His body under my hand was clammy with secretions. 

July 2, Tucson, Ari/. Rained in many places. Went about 5 to Sabmo Can- 
yon. At picnic spot two pools, the upper pool filled with R. pipiens. When al- 
most dark we went to the lower pool with its dam. At upper end we saw about 
12 dead females of Bit jo alvarius. Several large B. alvanus males jumped in. 
We continued to find males until we came to where Prof. Wehrle's small 
daughter had spotted a just perceptible gray spot in the shallow water (4-8 
inches). This proved to be a pair. There were six or eight pairs here. Brought 
in three pairs and put them in a tub. How the males did cluck! They sound 
like contented chickens or chickens when sleepy at night. At ti P.M. we found 
they had laid and we fixed some. What immense ropelike masses! They 
clucked when disturbed. 

July 3. This morning many B. alvanus eggs in the tub. One pair yet mated. 
The other pairs are broken. Went to Sabino Canyon. No B. alvanus to be 
found or eggs. 

Authorities' corner: 

J. G. Cooper, Calif., 1869, p. 480. A. I. and R. D. Oitenburger, Ariz., 1927, 

F. Mocquard, Gen., 1899, p. 168. p. 102. 

E. A. Mearns, Ariz., 1907, p. 113. F. W. King, Ariz., 1932, p. 175. 

A. G. Ruthven, Ariz., 1907, p. 506. C. F. Kauflfeld, Ariz., 1943, p. 343. 



TOADS: BUFONIDAE 



'39 






Plate XXIV. Bufo alvarius. 1,2,3,4. Fe- 
lales 



Plate XXV. Btifo americanus amcncanus. 
1,2,3,7. Males (X 1 /^)- 4- Male trilling while 
sitting in shallow water (X%). 5- Youn S 

(X 1 '-')- 6 - Coils of e ^ strin S s on bottom 
of a pond 



I 4 



HANDBOOK OF FROGS AND TOADS 



American Toad, Northern Toad, Hop Toad 

Bujo amencanus amcncanus Holbrook. Plate XXV; Maps 9, 10. 

Range: Manitoba eastward on latitude 50 N. to Gaspe and the Maritime 
Provinces. Minnesota south through Iowa to eastern Kansas and Oklahoma, 
thence eastward to northern Georgia, along Appalachians to central Virginia, 
thence to the coast and northward to Nova Scotia. A transition and Canadian 
zone form. Common throughout its range. 

Habitat: Common in gardens and cultivated fields, appearing more by 
night than day. During the sunshiny hours they seek cover beneath piazzas, 



B. a. cope? 




Map 9 



TOADS: BUFONIDAE 



141 



under board walks, flat stones, boards, logs, wood piles, or other cover. When 
cold weather comes, the toad digs backwards into its summer quarters or may 
choose another site for its hibernation. 

Size: Adults, 2^-4 Vi inches. Males, 54-85 mm. Females, 56-110 mm. 

General appearance: Short and fat in body, it has a short broad head and 
the snout is broadly circular. The lower surfaces are roughly granular, the 
back is covered with various-sized warts, some of which are large ones in pairs 
down the middle of the back. There arc three or four pairs of dark spots down 
the back, each with one large wart. The eyes are prominent. The arms and 
legs, hands and feet, are warty or roughly tubercular. There arc dark spots on 
the arms and legs, along the sides, and a few on the belly. Some males have 
yellow throats and considerable yellow on the underside of the base of the legs 
and in the groin. The general color is olive, with parotoids and crest brown. 

Color: Male. May 7, 1929. Back, sides, and tympanum dull citrine with 
olive-citrine or yellowish olive on hind legs and forelegs. Parotoids, crests, 
snout, and face ahead of eye huffy olive. Throat old gold, olivc-ocher, aniline 
yellow, or yellow ocher. Some of same color in axilla and on rear of brachium, 
also on groin and on lower belly. In latter region may be ochraccous-buff. 
Whole pectoral region with distinct scattered black spots; in fact these occur 
over entire venter except throat and center of rear belly. Lower throat oil- 
yellow or sulphine yellow. Some apricot yellow across arm insertion. Sides 
heavily spotted with large spots of black. On back is an obscure oblique black 
bar bordered with greenish yellow on upper eyelid. It is interrupted by the 
superciliary crest, only a small part showing inside this structure. One buffy 
olive tubercle in center of the eyelid is almost on the outer end of this bar. A 
more or less round black spot, greenish-yellow-edgcd, is just inside front end 
of long parotoid and just back of postorbital bar and end of superciliary crest, 
with tubercle in the center. Opposite the rear portion of parotoid are two 
oblique, posteriorly divaricating narrow bars with tubercles on forward end 
of each. Just beyond end of parotoids are the two largest round black spots 
with a large tubercle in each center. Halfway from parotoid to vent another 
such pair. Back of this pair arc suggestions of two or three obscure pairs. The 
tubercles in the five pairs are buffy olive. Femur and tibia are more or less 
crossbarrcd with black. Fore limbs spotted with black and a prominent 
oblique black spot across front of arm insertion. Rear of femur spotted or 
vermiculated with black and olive-citrine, old gold, olive-ocher, or primulinc 
yellow. Iris: dark in front of and behind pupil. Pupil rim citron yellow. Above 
pupil, mustard yellow and ochraceous-buff. 

Female. July 24, 1929. Color of back, light brownish or buffy olive; parotoids 
isabella or dark olive-buff; the bigger warts on the back, which arc in centers 
of dark spots, buffy brown; crests also buffy brown; stripe down middle of 
back, deep olive-buff, yellowish glaucous, or pale vinaceous-fawn, with more 
or less of the same color on oblique stripe that leads from parotoid to groin, 



142 HANDBOOK OF FROGS AND TOADS 

and also along the sides above and below the dark lateral band. The dark 
lateral band is brownish olive, olive, or deep olive. The tympanum may be 
buff olive or light grayish olive. The paired spots down the middle of the back 
are (i) oblique bars on upper eyelids; (2) spots inside the front of parotoid; 
(3) another pair of about the same size at rear end of parotoid with a spot 
near the middle line between these two. Back of these and farther from the 
middle line are (4) two larger, usually two-tuberclcd, spots sometimes with 
two small, single-tubcrcled spots inside each of these and nearer the meson. 
The next pair (5) are about the middle of the back and close to the meson. 
These arc the most prominent of the back and arc from i- to 4-tuberclcd. Then 
back some distance, close to the meson and slightly ahead of the vent, is an- 
other pair (6). These spots are black, fuscous black, or clove brown and are 
surrounded by reed yellow borders. Below the eye and tympanum is a black 
spot with a snuff brown center. The side of the face is clay color or huffy pink. 
The iris is same as in the male. Under parts are cream buff on the throat, be- 
coming primrose yellow on the belly and honey yellow or deep colonial buff 
on middle of lower belly and inner portion of buttocks. Among this is some 
orange-vmaceous or pale grayish vinaceous. In center of breast is a dark spot. 

Structure: Pnrotoids large and oblong, connected to the postorbital crest by 
a longitudinal ridge. Crests on the head form a right angle at the corner of 
the eye, one branch extending downward in front of the ear; males with dark 
throats; males with excrescences on inner-upper side of first two fingers and 
on inner carpal tubercle; many spiny warts, particularly on the hind limbs. 

Voice: "The note of the toad is long sustained, quite musical, rather high 
in pitch. Their low variable trills or pipings in chorus are rather pleasing and 
do not especially annoy, even though continued day and night during the 
height of their breeding season" (A. H. Wright, N.Y., 1914, p. 27). 

"In general it sounds like whistling, and at the same time pronouncing deep 
in the throat, bu-rr-r-r-r. It will be found that different toads have slightly dif- 
ferent voices, and the same one can vary the tone considerably, so that it is 
not so easy after all to distinguish the many batrachian solos and choruses on 
a spring or summer evening" (S. H. Gage, N.Y., 1904, p. 186). 

"The common toad of the mainland of New York State is called Bttfo amer- 
icanns. Its song is a sweet, trilling whistle, and may be imitated by whistling 
in a low monotone with drops of water held between the lips. Each individual 
song is prolonged for about thirty seconds. The prolonged song of the Ameri- 
can toad is a ready means of distinguishing it from the short song of the 
common toad (jowleri) of Long Island" (F. Overton, N.Y., 1914, p. 27). 

Breeding: They breed from April 5 to July 25, the crest about April 30. The 
eggs are in long spiral tubes of jelly, each egg %s-Me inch (1.0-1.4 mm.) in 
diameter; the inner tube %-]i2 inch (1.6-2.2 mm.), the outer tube %-% 
inch (3.4-4.0 mm.). The eggs, 4000-8000 in number, are laid in two strings, 
and hatch in 3-12 days. The small, dark, almost black tadpole i%2 inches (27 



TOADS: BUFONIDAF 143 

mm.) has an ovoid body broader near the vent than at the eyes. The dorsal 
crest is low, extending slightly onto the body, the tail short, its tip rounded. 
The tooth ridges are 73. After 50-65 days, the tadpoles transform June i to 
August at /4-Vis inch (7-12 mm.). 

Journal notes: June 8, 1911. Found at Crossroads Pond, Ithaca, N.Y., numer- 
ous Bufo americanus transformed and transforming, also several Rana syl- 
vatica transformed (at least 12 or 15). At Beebe Lake on south side near the 
spring are myriads of transformed Bujo americanus. 

June 9, Beebe, north side. On this side also arc countless Bujo americanus 
transformed. John Rich reports them numerous yesterday on Wait Avc. just 
over Triphammer bridge. Beebe, south side, Bufo transformed still numerous. 

June 12, Beebe, by the spring. Little toads are crossing the road above. They 
are not so numerous at water's edge. 

April 23, 1912, Crossroads Pond. Found two pairs of Bufo americanus lay- 
ing. Water cold, not much warmer than 4<)F. Found hidden males at varying 
distances from the pond on the hillside below. Many of them were not entirely 
under cover. They were in little pockets at the surface of the ground, their 
backs exposed and the skin as dark and dirtlikc as the dirt itself. 

Authorities 9 corner: 
C. C. Abbott, N.J., 1868, p. 805. 
A. II. Kirkland, Mass., 1897, p. 24. 
H. Carman, Ky., 1901, pp. 60-61. 

Hudson Bay Toad, Hudson Bay American Toad, Cope's Toad 

Bufo americanus copei Yarrow and Hcnshaw. Plate XXVI; Maps 9, 10. 

Range: Hudson Bay, James Bay. Gaige (Out., 1952, p. 134), who brought it 
out of synonymy, wrote: "During a vacation trip to the James Bay region in 
August of this year Mr. Calvin Goodrich kindly collected a scries of toads ( 10 
.specimens, ranging from 10 mm. to 69 mm. in length) for us at Moose Fac- 
tory." Riviere clcs Rapides, Labrador (F. G. Speck, Lab., 1925, pp. 5-6); 
Quebec. 

Habitat: "The colony . . . inhabits the abandoned clearing of a lumber 
operation of some years ago. Three or four decayed log stables and sheds and 
a surrounding mass of fine succulent meadow grass near the mouth of the 
river formed the environment. This small race is very active, and I have no- 
ticed that the living specimen spends much time buried about an inch deep in 
the sand . . ." (F. G. Speck, Lab., 1925, pp. 5-6). 

Size: Adults 1%-j inches (40-75 mm.). 

General appearance: "The brilliant coloration, the long, narrow parotoid 
glands, the greater width between the cranial crests and the fact that they are 
nearly parallel, are the most obvious characters. The smoothness of the ventral 
granulation, mentioned by Henshaw and Yarrow, is evident, but it may be 




*ri 

I 

o 

-Q 
-Q 
OQ 



I 

I 

u 



01 

-c 



CO 



3 
g 




OQ 





I 



TOADS: BUFONIDAE 145 

seasonal or due to preservation, and the hind limbs are shorter than is usual 
in Michigan specimens of B. amcricanns, the adpressed heel reaching the 
shoulder" (H. T. Gaige, Ont., 1952, p. 134). 

Color: Thirty miles cast of Moisie, Labrador, from H. W. Jackson, 1937. 
In captivity the male lived a year, the female two years. 

Female. The larger, more brightly colored of the two. The groin and rear 
axilla are brazil red. There is also a wash of the same color on rear under parts. 
A few dorsal tubercles along mid-back are claret brown or chestnut brown. 
The snout and parotoid glands are hazel. The middorsal stripe, light lateral 
stripe, and other light dorsal spots are pale smoke gray. The under surface is 
cartridge buff with heavy black mottling. The iris has glass green pupil rim 
and flecks of the same color. 

Male. The smaller toad is almost completely uniform in color. The snout, 
crests, parotoid glands, and warts are isabclla color. The back is olive- 
ocher or ecru-olive. The spots on rear of dorsum are yellowish olive, a few 
near the head are black. The belly is pale olive-buff, heavily spotted with black. 
The groin, rear of femur, and to a slight extent axilla are apricot orange. In 
the eye, the pupil rim is light orange-yellow with heavy flecking of the same 
in upper part of the iris. 

Structure: "Head subtriangular, broader than long; snout acuminate, pro- 
truding; head with well-marked groove, which extends to tip of snout; super- 
ciliary ridges strongly pronounced and terminating posteriorly in a slight 
knob; orbit bordered posteriorly by a similar ridge; upper jaw slightly emar- 
ginatcd; parotoids medium, elongated, twice as long as broad, perforated by 
numerous small pores, situated well back on the shoulders; not approximated 
to the tympanum, which is circular and large; limbs long and comparatively 
slender; palm rugose; a single well-developed tubercle; first, second, and 
fourth fingers about equal in length, the third longest; hind limbs rather 
longer than head and body together; tarsus and metatarsus with small and 
smooth tubercles; body above covered with small and somewhat roughened 
tubercles; under parts finely papillatcd; metatarsal shovel large" (H. C. Yar- 
row and H. W. Hcnshaw, Ncv., 1878, p. 207). 

Voice: Not of record. 

Breeding: Not of record. 

Journal notes: We have made notes on the Clausen, Trapido, and Vladykov 
material. Several explorers in Gaspe, Labrador, and Hudson Bay have re- 
marked : "What is the bright tood we saw ?" 

Authorities 9 corner: "Toads common in the valley of the Ste. Anne des 
Monts River, Gaspe Peninsula, differ strikingly from Bufo a. americanus. 
Comparison at the Canadian National Museum with material of B. ameri- 
canus copei from Moose Factory and from Cape Hope Islands in James Bay 
demonstrated that our toads were that subspecies. The coloration is much 
brighter than in B. a. americanus, with the black spots of the back fused on 



146 HANDBOOK OF FROGS AND TOADS 

the sides to form broken bands and blotches. These markings are usually so 
arranged that there is a broad light middorsal band, and frequently a light 
lateral stripe on each side, from the parotoid to the insertion of the hind limb. 
Our specimens also resemble B. a. copei in the pink coloration in the axils of 
the limbs and in the pink tipped warts over the dorsal body surface, as Cope 
(1895: 286) noted for a specimen from Moose River, British America, which 
he assigned to copei of Yarrow and Henshaw, although he did not consider 
that form valid. The parotoids not only are long and narrow, as Gaige (1932) 
has pointed out, but tend to be constricted towards their middle, while the 
chest and belly are speckled or in some individuals strongly mottled with 
black. 

"Sexual dimorphism is pronounced in a pair of living specimens which we 
have examined from Seal River, on the north shore of the St. Lawrence River. 
The female is brilliantly colored with black and pink dorsally, and with black 
marbling vcntrally, while the male is a drab yellow brown, with the black and 
pink much less pronounced and the middorsal stripe lacking. The underside 
of the male is spotted with black, though not so extensively as in the female. 
The parotoids of both male and female are narrow and long, while the cranial 
crests are well spaced. Our collection from the Ste. Anne River differ from this 
pair in that the males arc as brilliantly colored as the females" (H. Trapido 
and R. T. Clausen, Que., 1938, p. 120). 

"Bttfo americanus americanus (Holbrook). Common throughout the area 
and very abundant locally; usually found on land, but a few taken from wa- 
ter. Most of the toads were brilliantly colored, with patches of pink below and 
many small pink tipped warts on the sides. The dorsal spots were deep black, 
with several large warts in each. These toads thus tend to resemble B. ameri- 
canus copei Yarrow and Henshaw in coloration but lack the structural pecu- 
liarities of that subspecies. Logier (1928) has noted the more brilliant colora- 
tion of northern toads. The call was heard occasionally until July 17" 
(R. Grant, Que., 1941, p. 151). 

Northwestern Toad, Baird's Toad, Mountain Toad, Columbian Toad, Small- 
spaded Toad, Northern Toad, Western Toad 

Btifo boreas boreas (Baird and Girarcl). Plate XXVII; Maps 10, n. 

Range: Prince William Sound, southeastern Alaska, southeastward to east- 
ern Montana and northeastern Colorado and southwestern Utah, thence west 
to California (Mono County to northwestern California). 

Habitat: Terrestrial except at breeding time. Ruthven and Gaige (1915) 
stated the situation very well and tersely in their table of habitat distribution 
in Maggie Basin in northeastern Nevada. Bufo b. boreas occur in "general 
vicinity of larger water courses. Breed in water but can endure rather dry 
habitats during other times of the year." It was common in "basin floor valleys 



TOADS: BUFONIDAE 





Plate XXVI. Bufo amerlcanus copei 
- 1,2,3. Females. 4. Male. 



Plate XXVII. Bufo boreas boreas. 1,2. 
Males (X 1 /2). 3. Hind foot of male (X%). 
4,5. Females ( 



148 HANDBOOK OF FROGS AND TOADS 

and mountain canyons nearly to highest elevations. . . . Lives in tules about 
lake shores, along streams, and in mountain meadows" (J. Grinnell and C. L. 
Camp, Calif., 1917, p. 143). 

Size: Adults, 2^-5 inches. Males, 56-108 mm. Females, 60-125 mm. 

General appearance: This is a large brown, gray, or green toad with a light 
streak down the speckled back. The head is narrow and pointed in proportion 
to the fat broad body. There are rounded glands on the middle of the tibia. 
Lack of a discolored throat makes the males hard to distinguish from the 
females. 

Color: Aug. 31, 1929. Male. The light middorsal stripe is pale cendre green, 
pale dull green-yellow, or pale Veronese green, or even cartridge buff pr sea- 
foam yellow. The ground color along the back is dark olive or clove brown, 
becoming about one-half inch from middorsal line pale brownish drab, light 
drab, or drab-gray. As it approaches the belly it becomes an ecru-drab or pale 
drab-gray. The belly color is tilleul buff to pale vinaceous-fawn, or pale smoke 
gray. The throat, clear of spots, may be vinaceous-buff, tilleul buff, olive-buff, 
or pale smoke gray. Along either side of the middorsal line is an irregular row 
of warts. The next series outside begin to be conspicuously surrounded by 
black or black interspersed with citrine-drab areas. On the lower side the con- 
spicuous warts in the middle of the dark areas are lacking, and only the citrine- 
drab or water green spots are present. Across the belly these large black spots 
are very much in evidence as also on the inside of the hind legs out to the toes. 
The underside of the forelegs also have them, but they are localized on the 
rear and upper margins. On the rear of the belly and the underside of the 
buttocks the black spots are very fine, interspersed with cream color or light 
vinaceous-fawn. Here the tubercles are much larger and mainly cartridge buff 
or pinkish buff. The tubercles on the underside of the fore feet and tips of toes 
are light vinaceous-cinnamon. The parotoid is buffy olive or buffy brown, as 
is also the rim of the upper eyelid. The upper jaw and face ahead of the eye arc 
uniform grayish olive. There is a dark oblique area from the lower rear of the 
eye past the angle of the mouth. There is another dark area from the middle 
of the rear of the eye over the small tympanum and along the lower margin 
of the parotoid. There is a long longitudinal bar from the shoulder insertion 
along the front of the forearm. The eye is black or raw umber through the 
middle. The rim of the pupil is maize yellow. The lower pupil rim is of the 
same color or warm buff, but it is narrow and broken in its middle point. 
Above the upper pupil rim is the most conspicuous feature of the eye a prom- 
inent zinc orange or mikado orange longitudinal band. The lower part of iris 
is dotted with ochraceous-buff or zinc orange. 

Structure: No cranial crests; rounded gland in middle of tibia; body flat, 
broad; head narrow and pointed in proportion; no discolored throat in male; 
spread of hind foot usually more than 36 per cent of total body length. "Warts 



TOADS: BUFONIDAE 149 

on back show a tendency to run in longitudinal rows. Tibia with one large 
and one small parotoidlike wart located respectively in the central and the 
rear cross bars" (C. L. Patch, B.C., 1922, p. 77). 

Voice: Cope heard this form in a pond near the shore of Pyramid Lake, 
Nev. It was associated with a spadefoot (Scaphiopus intcrmontanus) of which 
Mr. Cope wrote: "Like other allied species, it was very noisy, almost obscuring 
the voice of the less vociferous Btifo" (E. D. Cope, Nev., 1884, p. 18). 

Our captive male toads, when held in hand and frequently at other times, 
give little birdlike chirps, very pleasant in tone. 

"At this time the males call with a high-pitched tremulous note, amplified 
by the vocal sacs distended beneath the chin" (G. C. Carl, B.C., 194?, p. 43). 

"The adults were large and tame, they usually walked instead of hopped, 
and when confined in a bag they scolded much like B. americanus" (A. G. 
Ruthven and H. T. Gaige, Nev., 1915, p. 14). 

Breeding: They breed from March to July. The eggs are in strings like those 
of B. boreas halophtlits of California. Several workers have seen the eggs but 
none has described them in detail (except Slater in manuscript). Our evidence 
is not sufficient, but it appears that the eggs are as follows: Two envelopes; 
no partitions between the individual eggs; eggs largely in a double row within 
the jelly tube; outer envelope large; 4.8-5.3 mm., some loose and distinct; 
inner envelope 3.5-3.8, distinct; vitellus 1.5-1.75 mm. 

The tadpoles are small, i Vi2 inches (27-44 mm ) with teeth %. There is no 
description of them on record. After a tadpole period of 30 to 45 days, they 
transform from July to September at %-Yi inch (9.5-12 mm.). 

"On the nights of June n and 12, 1928, these toads were seen in numbers 
in a larger pond on the beach at Kaslo [B.C. |. The males were calling and 
greatly outnumbered the females; nearly all the specimens collected or exam- 
ined at the pond were males. One male was seen on the beach in embrace with 
a dead female which was much dried and shrivelled. . . . 

"A female of 108 mm. taken at Kaslo on June n, 1928, had apparently fin- 
ished spawning; two other specimens of 81 and 101 mm. taken at Summcrland 
in July, one on the i7th, were full of eggs. A specimen of 93 mm. taken at Lyt- 
ton between the ist and 8th of July, 1925, had evidently spawned. 

"On the night of June 12, strings of eggs were found strewn among the 
grasses in the pond of water six or eight inches deep. The water temperature 
was 66 F." (E. B. S. Logier, B.C., 1932, p. 321). 

"Btifo boreas in Alaska: In the winter of 1896, Mr. A. W. Greeley, a student 
at Leland Stanford Junior University, gave me for examination 2 toads which 
he had 'taken swimming in a large lake near Prince William's Sound, Alaska, 
July 15, 1896.' . . . One of these specimens contains eggs which must have 
been nearly ready for laying" (J. Van Denburgh, Alas., 1898, p. 139). 

Several workers in Oregon, Utah, and Washington have found this form 



150 HANDBOOK OF FROGS AND TOADS 

breeding. James Slater has sent us some fine tadpole material but we are await- 
ing his Amphibia of Washington before venturing on further allusion to 
them. 

We have found on record no detailed description of the eggs or tadpoles. 

Journal notes: March 30, 1942. Went with James Slater to Sparaway Lake, 
Wash. As we approached the lake we saw a Bufo boreas borcas male. Caught 
it. 

April 7. East of Ashland and where road crosses creek (junction with Samp- 
son Creek) saw Bufo boreas boreas run over. 

April 8. Up creek above Trail, Ore. In one pool took a female and a male 
Bufo boreas boreas. 

Authorities' corner: 

E. D. Cope, Ore., 1883, p. 19. C. E. Burt, Gen., 19333, p. 351. 

M. M. Ellis and J. Henderson, Colo., C. F. Schonberger, Ore., 1945, p. 121. 

1915, p. 254. 

California Toad, Salt-Marsh Frog, Baird's Toad, Common Toad 

Bujo boreas halophilus (Baird and Girard). Plate XXVIII; Map 11. 

Range: According to Camp, the seven northern counties of California have 
Bufo b. boreas, while south of Eureka toads are Bufo b. halophilits. This spe- 
cies extends the length of the Great Valley and along the coast to northern 
Lower California. Somewhat east of Los Angeles, avoiding the southeastern 
deserts, the range cuts diagonally north of Owen's Lake to Lake Tahoc or 
Pyramid Lake in Nevada. 

Habitat: Open valleys, rarely wooded areas. In high mountains found in 
wet meadows and along lake shores. 

Size: Adults, 2? 3-4% inches. Males, 62-101 mm. Females, fx>-ii6 mm. 

General appearance: This is a large stocky toad with short limbs, green or 
greenish brown with a light streak down the back, and the back mottled with 
irregular dark areas which surround the warts singly or in irregular groups. 
The warts are rounded and the skin between the warts quite smooth. The 
white or yellowish under parts are in some individuals blotched with black. 

Color: April 22, 1928. Male. Stripe down the back is light lumiere green, or 
chrysolite green, or glass green. Upper parts mignonette green, light yellow- 
ish olive, or yellowish olive, becoming on the sides of the head buffy olive. 
Yew green or jade green on sides of body with background pale glass green or 
glass green. Under parts white. Buttocks buffy olive or buffy brown. A few 
dark spots on under parts. Throat not different from breast. Iris chrysolite 
green above and below with black; black through the middle of the eye. Iris 
rim cream-buff. 

Another male. Sides and upper parts dark olive-buff. Upper parts citrine 
becoming sulphine yellow, olive lake, or ecru-olive, then reed yellow or prim- 



TOADS: BUFONIDAE 



I j I j 1 1 B.b. halophilus 

III 




Map n 

rose yellow, then cartridge buff on lower side. Some warts with honey yellow 
on hind legs and sides; browning olive on warts along median line. Median 
line reed yellow from snout to vent. Under parts pale olive-buff. Throat with 
wash of cream-buff. Buttocks and lower belly Isabella color and purplish. 
Black areas of upper parts with buckthorn brown centers. A few spots of 
ochraceous-orange along sides. 



152 HANDBOOK OF FROGS AND TOADS 

Female. Grayish olive above; upper parts of fore and hind legs light grayish 
olive; or olive-buff upper parts; or light mineral gray on all upper parts. Stripe 
down back chalcedony yellow or citron yellow. Upper parts with numerous 
black areas; these often with olive-buff centers. Along either side of median 
line are warts, some with dark olive-buff or picric yellow centers. Throat olive- 
buff, pectoral region and upper belly pale olive-buff. Under part of femur and 
lower belly olive-gray or deep olive-gray. Iris, sulphur yellow pupil rim, above 
which are black reticulations, then sulphur yellow becoming pale yellow- 
green, then beryl green rim on outside above, but seafoam yellow or ivory 
yellow behind and in front. Lower part of iris black with a little cream-buff or 
pinkish buff. 

Structure: No cranial crests (except occasionally in very large individuals) ; 
parotoids elongate, widely separated; two long metatarsal tubercles, inner 
with a free blunt end; spread of hind foot usually less than 36 per cent of head 
and body length; glands on tibia present; no external vocal sac apparent in the 
males; interorbital space only slightly greater than internasal space; first and 
second fingers equal; not so heavily pigmented as B. boreas boreas. 

Voice: Its song is a slow, deep-toned, prolonged trill. 

"The males, while in the water, utter a series of low mellow tremulous notes. 
In chorus the notes may be compared to the voicings of a brood of young do- 
mestic goslings. The call of each male is uttered for a second or two and re- 
peated at short intervals, so that a practically continuous chorus issues from 
a breeding colony. To human ears the notes lack carrying power and can 
scarcely be thought to be of use in attracting toads at any great distance from 
the pools where the males are calling. Occasionally males in their daytime 
retreats will utter the notes once or twice. This species does not have an en- 
larged vocal pouch such as is possessed by many species of toads (for example, 
americanus, cognatus, tvoodhousii)^ and this lack of a 'resonating pouch* is 
probably responsible for the small volume of sound uttered. Calling, with 
halophilus, is to be heard in the daytime as well as at night" (T. I. Storer, 
Calif., 1925, p. 177). 

Breeding: This species breeds from January to July, according to the climate 
of location. The eggs are in long strings laid at the margins of ponds or at 
edges of flowing streams and are occasionally in two or three rows. There are 
no partitions between the eggs. The vitellus is Mo inch (1.7 mm.), the outer 
tube % inch (5.0 mm.), the inner tube }4 inch (3.6 mm.). The dull blackish 
medium tadpoles are 2% inches (55 mm.). The tooth ridges are %. After 28 
to 45 days, the tadpoles transform from April to August at %-% inch (12-15 
mm.). 

Journal notes: Feb. 12, 1942. Visited W. M. Ingram. He had a pair of B. bo- 
reas halophilus mated axillary fashion. 

March 6. About 10 A.M. Anna and I went to Stanford artificial lake to look 
for A. t. calif orniense. Didn't find any. At last found at end of a pond a mated 



TOADS: BUFONIDAE 



'53 




Plate XXVIU. Bufo boreas halophiltts Plate XXIX. Bufo boreas nelsoni (X%). 

. i. Male. 2,3. Females. 1,2,3,6. Males. 4,5. Females. 



154 HANDBOOK OF FROGS AND TOADS 

pair of Bufo b. halophtlus. The male has a friendly chuckle, which it gives at 
times while mated. Presently I came to an area in the water near a fallen tree. 
Here, all about, at least 10 feet out from the bank in water 12 inches to 2 feet 
deep, were long strings of eggs. The pairs must be very restless. The strings 
are wound around many tussocks and are often in two bands for a long dis- 
tance. The spider web is quite different from our usual Bufo a. amcricanus 
bands, which are in the very shallow muddy edges and very heavily massed* 
The egg file is a double row of eggs and somewhat like that of Bufo joivleri 
and Bufo woodhottsu. Later near edge saw a pair "scrunch" down under the 
edge of a board and beside a tussock. Captured them and another single male. 

April u, around Chico, Calif. In the forenoon went out beyond golf links. 
Found toad eggs. Later in grass, beside a reservoir took a toad. In edge of a 
stream saw a toad "scrunched" down in grass. Caught it. In pools around 
reservoir no end of tadpoles. Some Bitfo just hatched. 

Authorities 9 corner: 
H. C. Yarrow, Nev., 1878, p. 208. 
J. Grinnell and T. I. Storer, Calif., 1924, pp. 655-656. 
J. R. Slevin, B.C., 1928, pp. 95-96. 

Amargosa Toad 

Bufo boreas nelsom Stejneger. Plate XXIX; Map n. 

Range: Southern and eastern Nye County and northern Lincoln County, 
Nev., to Resting Springs, Morans, and Lone Pine, Owen's Valley, Calif. 

Habitat: "So far known from three separated localities, but most character- 
istic population is in the upper part of the Amargosa River. Apparently this 
toad is more closely restricted to water than even its near relatives which in- 
habit more humid districts" (J. M. Linsdalc, Nev., 1940, p. 204). 

Size: Adults i%~3% inches. Males 42-68 mm. Females 46-89 mm. 

General appearance: "Similar to B. boreas: skin between warts smooth; 
snout protracted, pointed in profile; webs of hind legs very large; soles rather 
smooth; limbs shorter, elbows and knees not meeting when adpresscd to the 
sides of the body; inner metacarpal tubercle usually very large" (L. Stejneger, 
Calif, and Nev., 1893, p. 220). 

Color: Springdale, Nev., May 13, 1942. This is another boreas type toad. 
The general background of back is buffy olive, with the side of face and upper 
part of hind limbs the same. The stripe down mid-back is primrose yellow to 
pale olive-buff. The parotoids bear the most prominent color of the back, being 
tawny-olive to tawny. There are three paired rows of tubercles: one along 
either side of middorsum, one from parotoid backward, and the third an ir- 
regular intermediate one. All these tubercles have centers of cinnamon-brown 
or mikado brown. Back of the angle of mouth is a patch of ochraceous-tawny 
or ochraceous-orange, which color extends backward topping a row of very 



TOADS: BUFONIDAE 155 

flat tubercles. Below these the black spots become large and prominent s each 
centered with a light-tipped tubercle. There is a prominent black area below 
the eye, another from tympanum downward, and sometimes one below the 
nostril, all three along the upper labial border. The upper eyelid bears some 
tawny-olive or tawny. The iris is black with a cream color or citron yellow 
pupil rim, the same colors flecking the whole eye. There is a prominent 
oblique bar of primrose yellow or marguerite yellow at the front edge of eye. 
The throat is primrose yellow to marguerite yellow almost clear of spots; the 
rest of under parts arc pale olive with scattered black spots. 

Males and females are alike in color, but after being in captivity and after 
breaking their axillary mating hold the males were very black and the female 
markedly lighter, this past night. When first brought in all five toads looked 
quite light. This afternoon they are much the same in color pattern. 

Structure: "The 51 specimens now on hand from Oasis Valley and the 
Amargosa River near Beatty . . . contrast with toads of the species Bnjo boreas 
to the north and west in the following characters: Small size: the largest adult 
in the lot measures 72.5 mm. in head and body length. This is a little more 
than half the maximum size of Bufo b. boreas. The narrow, wedge-shaped 
head, especially when viewed from below, is one of m the striking peculiarities 
of this toad. This seems to be correlated with the snout protracted and pointed 
in profile as mentioned by Stejneger. Limbs so reduced that when adpressed 
to sides of body elbows and knees do not meet, as was pointed out by Stejneger. 
Small feet and reduced webbing are especially noticeable and are just the re- 
verse of Stejneger's statement, which may not have been as he intended. Re- 
duced spots below in all the large individuals contrasts with boreas and with 
all small ones from the Amargosa Valley. Possibly the old ones tend to lose 
ventral spotting. Smooth skin and small weakly developed warts which char- 
acterize this lot may be an indication of close restriction of these toads to the 
water. Inner meiacarpal tubercle appears to average large, as Stejneger indi- 
cated" (J. M. Linsdalc, Ncv., 1940, p. 204) . 

Voice: Beatty, Nev. The male frequently gives a short cheerful chuckle. We 
don't know that we can tell this from the chuckle of B. exul. Both are boreas 
types. 

Breeding: J. M. Linsdale, D. M. Hatficld, and J. K. Doutt secured trans- 
forming toads in May from 15-18 mm. in length. 

"Near Beatty, about May 20, 1931, many small young toads were picked up 
near the stream; large tadpoles were obtained here on May 4, 1936" (J. M. 
Linsdale, Nev., 1940, p. 204). 

Journal notes: May 13, 1942. At Springdale, Nev., in the hamlet itself, saw a 
muddy pool with full-grown tadpoles; looked around but saw no toads. Came 
to Beatty about 4 P.M., walked i% miles north along Amargosa River. No 
toads. Went to Springdale. In her pool, Mrs. Howard said they had tadpoles. 
I looked at them. They were about one-third grown. She said the eggs were 



156 HANDBOOK OF FROGS AND TOADS 

laid two weeks ago in the water-lily box. "They were in strings with a little 
egg, then space, then another egg." One man said these toads could be seen 
almost any night. 

Last night after 8 we went from Beatty, Nev., to Springdale. With dogs 
barking and peacocks calling we thought our mission ruined. We went to 
our tadpole pool. Across the pool I saw a light toad or frog. Made for it and 
caught my first Bufo b. nelsoni, the Amargosa toad. Soon saw another resting 
in shallow water. Went to another pond near by and here soon espied two 
more a large male and a small female. None are mated. These are shallow 
pools, mucky, and no more than 2 to 8 inches deep. Returned to the first pool 
and here another male was suspended in water in the weeds. These are not 
hard to capture. Where were they in the daytime when I was here? Last night 
about 5, at the suggestion of Mrs. Howard, we looked up Bobbie Shoshone, 
an alert Indian boy, and told him what we wanted. 

May 14. Bobbie Shoshone went out last night and caught one Bufo b. nel- 
soni and several Hyla regtlla. This morning he went out again and secured 
three more toads. 

Authorities 9 corner: 

L. Stejneger, Calif, and Nev., 1893, pp. 220-221. 
J. M. Linsdale, Nev., 1940, p. 204. 

Southern California Toad, Arroyo Toad 

Bufo calif ormcus (Camp). Plate XXX; Map 8. 

Range: Coastal region of southern California San Luis Obispo Co. (Miller 
and Miller, 1936) through Ventura Co. (Camp, 1915) and San Diego Co. 
(Klauber, 1928) to lower California on Rio Santo Domingo (Klauber, 1955; 
Tevis, 1944). 

Habitat: Arroyo stream beds; inland valleys and foothills. 

"In San Diego County the range seems largely restricted to the sandy 
washes of the rivers in the Upper Sonoran Zone" (L. M. Klauber, Calif., 1931, 
p. 141). 

"The river bottom has marginal growths of oaks and cottonwoods on sandy 
beaches with willow and Bacchans thickets bordering the stream. The small 
flow of clear water gave promise of continuing well through the summer. The 
stream subdivided frequently and at places formed comparatively quiet pools 
in the gravel as much as 18 inches deep. It was within a few feet of one of 
these that a trilling animal was watched in the beam of a flashlight as it sat 
half submerged in a puddle not more than 2 feet across. The fore part of the 
throat was moderately distended, not to compare with the Hyla rcgilla which 
were calling on all sides" (L. and A. H. Miller, Calif., 1936, p. 176). 

Size: Adults, i%-2% inches. Males, 44-58 mm. Females, 44-68 mm. 

General appearance: This is like a small Great Plains toad (Bufo cognatus) 



TOADS: BUFONIDAE 



157 



but is more uniform in color on the back. This little toad is very short and 
thick in body, very short in head, and the arms and legs appear very stolid. 
The foot is a little longer than in B. cognatus. The eyelids and parotoids are 
tuberculate. Several of the larger warts have spiny tips, in some cases several 
tips to one wart. In a toad that has recently shed its skin, there is a row of con- 
spicuous light tubercles on the side, bordered above and below by irregular 
black lines. In the young the light and dark areas of the head and back are 
strongly contrasted. Since 1930 we have seen most of the Bufo californicus 
specimens in collections. Those who know it best wonder if it should be com- 
pared to Bufo compactilis rather than Bufo cognatus. Somehow we cannot 
now bring ourselves to place woodhoitsii, w. fowleri, or californicus in the 
lower f Ar00Mappet group of Bufo compactilis and Bufo cognatus. 

"A toad with divergent head crests, a nasal boss, short slightly divergent 
parotoids and with an internal, cutting tubercle on hind foot; femur short as 
in Bufo cognatus cognatus, the Great Plains toad. Size medium; parotoids 
wide; coloration nearly uniform, without large spots; no vertebral streak; ex- 
ternal tubercle on hind foot small and rounded, not provided with a cutting 
edge" (C. L. Camp, Calif., 1915, p. 3?i). 

Color: Females. Warner's Hot Springs, San Luis Rey River, Calif., from 
L. M. Klauber, described June 12, 1929. Upper parts olive-brown or dark olive. 
On top of snout are two parallel black bars to level of front of eye where they 
connect with transverse bar. From middle of upper eyelid an oblique back- 
ward bar of black extends inward but does not meet its fellow. Just inside the 
front end of parotoid is faint black spot centered with a wood brown or avel- 
lancous wart. Opposite back end of parotoid two narrow black spots meet at 
an angle pointed forward. Just back of each of these and farther from meson 
is a larger round spot. About midway from eye to vent is a pair of spots near 
the meson. Thereafter to vent spots are very small, possibly three or four either 
side of meson. Ahead of oblique black bar of eyelid is a prominent light band 
across top of head, olive-buff in young or wood brown to avellaneous in adult. 
Same color on front third of parotoid, between angle spots and back of next 
pair of spots to each side of meson midway from vent to eye. On each side an 
oblique line of olive-buff warts extends to the groin with black line below it 
and sometimes an avellaneous band above with a black line above that. Sides 
olive-buff or pale olive-buff. Under parts white. Throat pale olive-buff or til- 
leul buff. Underside of feet and forefeet vinaceous buff. Underside of hind 
legs and rear belly light vinaceous fawn or light grayish vinaceous. Two or 
three olive-buff warts back of angle of mouth. Two black crossbars on fore- 
arm, none on brachium. One-half crossband on front of femur and two on 
rear half of tibia. Each bar has centers or warts of avellaneous or wood brown. 
Space between tibial crossbars olive-buff or pale olive-buff. Dark spot below 
eye; oblique olive-buff area in front of eye in young, wood brown in adults. 
Iris black. Pupil above and below, but not back or front, with prominent rim 



158 HANDBOOK OF FROGS AND TOADS 

of light or pale chalcedony yellow and with spots of this color or apricot buff 
on iris. 

Male. Glen Lonely, San Diego Co., Calif., from L. M. Klauber and L. Cook, 
March 28, 1930. The two males in hand have the light band between eyes, deep 
olive-buff or yellowish citrine. The front third of parotoids is not conspicu- 
ously different from the rest of parotoid as in female described. It may have 
touch of vinaceous buff. The upper parts of these males are dull citrine, gray- 
ish olive, or dark grayish olive. The prominent warts are tipped with one to 
several spines. Sometimes, but not always, these are dark or brownish tipped. 
The dorsal spots of these males are not so prominent as in females described. 
The short wood brown or avellaneous band down mid-back is inconspicuous 
in these males. The lateral oblique line of olive-buff warts is not conspicuous 
in these males nor in the female in hand. (Another female with freshly shed 
skin is much brighter in color and this line of light warts appears.) The throat 
is slightly different from the rest of under parts. The middle and lower throat 
is vinaceous-ibuff. On upper jaw below the rear of eye, in both males and 
females, is a light vertical band of wood brown, avellaneous, or vinaccous-buff. 
Iris of each male is more spotted and not so black as in the female in hand. 

Structure: Parotoids wider and longer (width in length 1.75-2.0) than in 
B. cognatus (width in length 1.5-1.8) ; parotoid interval wide, twice the gland 
width. Occasionally a postorbital bar present and the posterior end of the cra- 
nial valley slightly suggesting embossment. One cutting metatarsal tubercle 
and one small one; femur short; foot about one-half webbed; hind limbs long; 
half or moie of femur free from body skm. 

Description of type: "Size medium; hind legs very short, femur almost en- 
tirely enclosed in skin of abdomen; head short and thick; nasal region elevated 
into a bony protuberance; longitudinal cranial crests more or less united across 
median region, and slightly divergent; transverse crests divided by width of 
median groove; parotoids oval, slightly divergent and very broad; inner tu- 
bercle of hind foot with a sharp edge; outer tubercle very small, rounded and 
without cutting edge; eyelids and back evenly tuberculated; tympanum oval, 
shorter than diameter of eye; a dozen or more large whitish tubercles below 
and just posterior to the tympanum" (C. L. Camp, Calif., 1915, p. 332). 

Voice: "Three animals were trilling, spaced about equally along a quarter- 
mile of stream bed. . . . There is considerable resemblance to the louder, 
more raucous call of cognatus, even granting the difference which Myers em- 
phasizes" (L. and A. H. Miller, Calif., 1936, p. 176). 

"The call of cognatus is a trilled rattle, with much of the timbre of Acns in 
it. The call of calif ornictts is a sweet trill reminding one of B. americanus but 
somewhat lower and less prolonged" (G. S. Myers, Calif., 1930, pp. 74-75). 

"The call is a beautiful, penetrating trill with a peculiar ventriloquistic qual- 
ity which makes it difficult to determine the direction from which it ema- 
nates" (L. M. Klauber, Calif., 1934, p. 6). 



*59 

Breeding: May and June. 

We need more descriptions of the egg masses, individual eggs, tadpoles, and 
transformation. 

Journal notes: March 25, 1930. Our experiences with these toads in life have 
been entirely with material sent us by L. M. Klauber. At present we have four, 
two males and two females in our toad garden. During the day they keep 
themselves buried in the moist soil with just their heads out. They are not 
always sleeping, however, as they promptly stir if an ant is thrown near them. 
Toward evening they are more active and hop around. One toad seems to 
have adopted an evening perch in a pot of herb robert. During the evenings 
the males sometimes give brief calls, just enough to suggest their little trill. 

April 24, 1942, Atascadero, Calif. Went to Salinas River northeast of Santa 
Margerita for toads, hoping for Bufo californicus. No luck! 

April 25. Big Pines Area: a public camp just beyond (E.) of Jackson Lake. 
Came to shallow pool beside the road, shadowed with willows. The outlet was 
through grassy field and row of willows and shortly fell rapidly into a deep 
cut. While Bert explored the willow tangle outlet, I walked slowly around 
edge of pond. There were tussocks of grass. I soon came to a shallow corner 
and there tangled on surfaces of grasses were two toad egg strings. These 
seemed small and narrow for Bufo b. halophilus. Eggs close set and tending 
to be in a double row, type like Bufo woodhousii or Bufo fowleri. Are these 
Bufo californicus or Bufo b. halophilus? Not the same as Bufo b. halophilus 
we saw at Palo Alto. If they are Bufo californicus eggs they are totally unlike 
Bufo compactilis or Bufo cognatus eggs. 

May 5, Have we arrived at Klauber's "3% or 4 miles south of Buckman 
Springs" where he caught Bufo calif ormcus, i.e., at the bridge below the con- 
fluence of La Posta and Cotton wood Creek ? Here is a wide expanse of sand, 
willow along creek, oaks farther back. We took here three T. o. hammondii, 
two H. regilla, and one R. a. draytonii. This would be an ideal place to get the 
toad if it were out. Saw no Bufo, Rana, or Hyla regilla tadpoles. 

May 6. Collected at Green Valley Falls Public Camp on Swcetwater River 
(creek). Just above water crossing, found two fresh complements of Bufo. 
Each is a small complement of a small toad. They are more or less in double 
arrangement in tube like Bufo woodhousii, Bufo w. fowleri, Bufo b. boreas, 
and Bufo b. halophilus. Two other complements are hatched. 

Robert Livezey of Stockton, Calif., has examined these eggs. He concludes 
that they are different from Bufo boreas halophilus and from the Bufo boreas 
boreas he knew at Corvallis, Ore. We both suspect they are Bufo californicus, 
the form we so often sought in vain. Our summary follows: "Files or strings 
with a continuous gelatinous encasing. One envelope present. Envelope more 
than 5 mm. in diameter; 5.62-6.12 mm., average 5.77 mm.; distinct and rela- 
tively firm. Vitcllus 1.25-1.62 mm., average 1.42 mm.; black above and gray 
below. Average of 42 eggs per inch; several thousand per female. Deposited 



160 HANDBOOK OF FROGS AND TOADS 

in tangled strings on bottom of pool among leaves, sticks, gravel, mud, etc. 
Range, coastal area of southern California from upper Salinas Valley to Lower 
California on Rio San Domingo. Season May and June. Bufo californicus" 
We hold that Bufo cognatus, B. b. boreas, and Bufo b. halophilus have two 
envelopes in a file. Our identification of these eggs as B. californicus is pre- 
sumptive and circumstantial. 

Authorities 9 corner: 
C. L. Camp, Calif., 1915, p. 332. 
G. S. Myers, Calif., 1930, pp. 75-76. 
L. Tevis, Jr., L.C., 1944, p. 6. 

Yosemite Toad, Yosemite Park Toad 

Bufo canortts Camp. Plate XXXI; Map n. 

Range: Yosemite National Park, high central Sierra Nevada, 7,000-11,000 
feet altitude. West branch of McGee Creek trail, Mono Co., Calif., at 10,500 
feet, Sept. 18, 1939 (O. R. Smith and K. Stanton). Two stations in Tuolomne 
County, 35 miles northwest from Tioga Lake locality (I. L. Wiggins, Calif., 
1943, p. 197). Occupies the Canadian and Hudsonian life zones, extending 
even into Alpine-Arctic (J. Grinnell and C. L. Camp, Calif., 1917, pp. 143- 
144) . (See Map i, p. 6.) 

Habitat: Wet mountain meadows, margins of streams and lakes. 

Size: Adults, r%-3 inches. Males, 46-64 mm. Females, 45-75 mm. 

General appearance: This small toad has a moist skin and the skin is smooth 
between the few large, low-rounded warts. The large broad parotoids marked 
with russet pattern or crossed by the pattern of the back and the glands on 
legs are the conspicuous marks. This species is dimorphic, the female with 
conspicuous touches of black, yellow, and russet, the male somber olive. 

Color: Male. Described Oct. 12, 1929. The male lacks the striking pattern 
of the female, being an almost uniform yellowish olive to olivaceous black 
with a broken thread of yellow down midback. The black-dotted borders of 
the darker markings and dark areas around warts are evident to greater or 
less degree as the toad is lighter or darker in color. These may be outlined by 
broken threads of yellow a delicate tracery. Along the sides where belly 
merges into side may be a yellowish band. 

Female. Yosemite Park, Calif., from C. A. Harwell and Dr. H. C. Bryant. 
Beginning with upper eyelid there extends along the back either side of the 
narrow vertebral line a longitudinal band of black. In this there are from time 
to time warts, dark olive-buff or isabella color. On rest of back are large spots 
of black with these colored warts in them. On sides, large black spots, but with 
few or no warts in them. Middorsal stripe may be pale lemon yellow to sul- 
phur yellow or primrose yellow. Background color is lime green, citron green, 
light chalcedony yellow, sulphine yellow, or olive-lake. Back of angle of mouth 



TOADS: BUFONIDAE 



161 




Plate XXX. Bufo californicus (X%). 1,2. Plate XXXI. Bufo canorus (X%). 1,2,5. 

Males. 3,4. Young. 5. Female. Males. 3,4. Females. 



162 HANDBOOK OF FROGS AND TOADS 

is a xanthine orange wart. Along on sides and base of arm insertion are some 
mars yellow or xanthine orange spots on ground color. Fore limb heavily 
spotted with black. Hind limb to tarsus same, also on rear of femur. Edge of 
upper eyelid and rear part of parotoid xanthine orange or less bright. From 
upper eyelid across parotoid is a longitudinal band of pale drab gray. The yel- 
lows of the background color encircle the black spots and in alcohol or formal- 
dehyde make them appear white encircled. Under parts white or pale smoke 
gray or pale olive gray. Few spots on side of lower belly, and a very few on 
rear of buttocks or on ventral side. All in all this specimen is pure color below. 
Iris with background color plus xanthine orange or mars yellow; this reticu- 
lated with black. 

Half-grown specimen. Oct. 12, 1929. Unlike large female, under parts are 
ivory yellow or white with spotting from angle of mouth across pectoral re- 
gion to angle of mouth; also on sides of belly and somewhat on belly proper. 
Buttocks and lower belly light mouse gray. Back darker; no middorsal stripe 
or slight. Spots more numerous. Background color drab or light drab except 
where surrounding black spots, then cartridge bull or sulphur yellow. There 
is hardly any impression of yellow in upper part as in full-grown female. Paro- 
toid with prominent antique brown or "rusty-like" spot. Same wart back of 
angle of mouth. Little touch of same color around vent. Sides of head and face 
spotted, as is back. Iris black with some sulphur yellow, pale brownish vina- 
ceous, or some "rusty," but more black than full-grown adult female. 

Male. Percgoy Meadows, Yosemitc National Park, Calif., from Carl E. Van 
Deman and G. B. Upton, June, 1930. These males vary greatly in color when 
kept in tub of dirt and when exposed to sunlight. The back may be olivaceous 
black or dark olive, becoming deep olive on the sides. The wart at corner of 
mouth may bear a dash of ochraceous-orange or orange-rufous and the paro- 
toids a wash of Sanford's brown. There is a reed yellow thread down the back, 
broken and irregular. Lower surfaces are white to seafoam yellow with many 
black dashes around edge of jaw, in pectoral region, and on belly. The iris 
bordering pupil is bull-yellow, then a ring crossed with network of black lines 
gives medal bronze appearance. Along the upper edge is a band of kildare or 
glass green specks. 

Held in hand for this description, the color has grown lighter, the back is 
yellowish olive, the sides light yellowish olive. The bars on legs appear as 
dark greenish olive bands with black dots around them forming irregular bor- 
ders. Some dark greenish olive areas appear on back surrounding some of the 
warts. Some of the warts have wash of Sanford's brown or amber brown. 
There is a dash of reed yellow in front of eye. After photographing, the back 
had become yellowish oil green. 

Structure: Short, broad, raised parotoids close together; muzzle rounded; 
cranial crests lacking or slight in some males; legs short; gland on hind leg; 
interorbital space narrow; internasal space wide; warts few, low, flattened, and 



TOADS: BUFONIDAE 163 

rounded; foot with two metatarsal tubercles; skin moist like that of a Ran a. 

This toad has a distinct rounded ridge from the nostril over the outer edge 
of the eyelid. This gives the canthal region a distinct border above and with 
the steep snout gives the toad a square-snouted appearance. The head is not 
so short, however, as in Bufo cognatus. The nostrils are widely separated. In 
general shape, the toad reminds us of Bufo boreas halophilus. Its color is inter- 
mediate between the color of that form and the heavy inky blotches and con- 
trasting light areas of Bufo cognatus. 

"In profile, lack of head crests, small tympanum, and short legs this toad 
resembles Bufo boreas and its subspecies, but may be distinguished at once 
from these forms by its smaller size, enormous width of parotoids, slight in- 
terval between parotoids, very smooth skin, absence of a broad vertebral stripe, 
and markedly different color pattern in both sexes. In extent of webbing of 
hind foot the present species most nearly resembles B. boreas halophilus 9 its 
near neighbor in the southern Sierra Nevada and the San Joaquin Valley. 
Specimens of B. boreas boreas from Mono County, directly to the east of the 
range of canorus, have the large hind foot characteristic of the more northern 
subspecies" (C. L. Camp, Calif., 19163, p. 60). 

Voice: The song is a sustained, rapid, melodious trill. 

"The specific name selected, canorus, refers to the long-sustained, melodi- 
ous trill uttered by this toad. This diurnal singing accompanies the breeding 
activities, which take place as soon as the snow melts from the Sierran mead- 
ows, June i to July 15. Many of the females captured at this time contained 
mature eggs" (same, p. 62). 

"The mating song of the Yosemite toad is a sustained series of ten to twenty 
or more rapidly utteied notes, constituting a 'trill/ and the whole song is re- 
peated at frequent intervals. The notes, though mellow in character, carry 
well considering the size of the animals, and have a ventriloquial quality 
which makes it difficult to locate any one animal by sound alone. When a 
number of males are giving their songs m the same place the songs overlap 
one another so that the general chorus is continuous. There is some difference 
in the pitch at which the several members of a group sing, varying perhaps 
with the size of the individual toad. . . . The notes recall the courting song 
of the Texas nighthawk" (J. Gnnnell and T. I. Storer, 1924, p. 659). 

"The toad chorus at different levels may begin at least as early as May 20 
and last until July 9; and according to our experience, singing is carried on 
quite through the daylight hours and into the early evening at least" (J. Grin- 
nell and T. I. Storer, Calif., 1924, p. 660). 

Breeding: These toads breed during late spring and summer, May 20 to 
July 16. G. S. Myers at Peregoy Meadows June 20, 1928, secured transformed 
toads 10% nim. long and tadpoles 27 mm. long with teeth %. |T. I. Storer, 
Calif., 1925, p. 42, figures teeth %.] 

Journal notes: "J une 5> I 93 ? Peregoy Meadows. Saw at least two dozen old 



164 HANDBOOK OF FROGS AND TOADS 

and young. Majority were young. They were not so abundant as the croaking 
tree frogs, Hyla regilla. Hadn't distinguished sexual dimorphism until I 
found a mated pair in a very wet grassy meadow, water three or four inches 
deep when one steps in it. These meadows are surrounded by snow fields. 
Found no toad eggs. Toads must have just come out" (C. E. Van Deman). 

July 24, 1934. Between Tuolomne and Dana Meadows. An intermittent 
stream went through the meadow leaving many isolated pools. No standing 
pools over the whole surface, though the water comes almost to the surface 
in some of the "lat" holes. Our first sight of Bufo canorus was of something 
scuttling along the runway of a "rat" hole at the base of a shrub. We found 
two small toads here. Near the pools we found many transformed ones, and 
several adults were in holes or under overhanging turf at the edge of the pool. 
We found one adult under a stump. Under a shrub were two in water and 
one on land. Many tadpoles and transforming toads were in one of the rather 
shallow pools; many transforming ones just under the edge of tussocks in the 
water. 

At Dana Meadows. Occasionally in a pool with sedges or vegetation or in a 
more marshy pond, Bufo canorus adults started to swim out into the water. 
In a few pools may have seen as many as 20 to 25 Bufo canorus tadpoles. The 
predominant tadpoles are of Hyla regilla. 

Authorities' corner: 
C. L. Camp, Calif., 19163, p. 61. 
J. Grinnell and T. I. Storer, Calif., 1924, pp. 658-659. 

Great Plains Toad, Say's Toad, Western Toad, Plains Toad, Texas Toad, 

Western Plains Toad 

Bufo cognatus (Say). Plate XXXII; Map 10. 

Range: Imperial Valley, Calif., up Colorado River to southeast Nevada, 
through Salt Lake, Wyoming, Montana to Alberta, thence to extreme western 
Minnesota and Iowa, northwest Missouri, western two-thirds of Kansas and 
Oklahoma southward to Texas coast; also in Mexico. 

Habitat: Grazing lands or agricultural lands of the Great Plains, along 
irrigating ditches, flood plains of streams, and overflow bottom lands. 

Size: Adults, i%-4 inches. Males, 47-95 mm. Females, 60-99 mm - 

General appearance: This is a large broad-bodied toad with a general color 
of brown, gray, or greenish. There is a light middorsal stripe, and the back is 
marked with dark spots. These spots may be broken up and the light areas or 
borders become more conspicuous so that the toad appears obliquely streaked 
with light bars on the sides. The spotted condition seems the more common. 
Often there are four pairs of bright green spots down the back, the pair at the 
rump leading diagonally to the groin. There are green spots on the legs, and 
an oblique row of green spots extending backward from the sharply raised 



TOADS: BUFONIDAE 165 

parotoid. There is a green band along the side. These green spots are partially 
outlined with black on the green and cream on the outer edges. The under 
parts are light, including the throat. In these males the light throat extends 
back as a flap over the thin, dark-colored skin of the lower throat, which ex- 
tends forward when inflated. In another toad the green spots are more broken 
up, and the green band on the side becomes broken into spots. The hands and 
feet are light with dark tips. 

Color: Male. Bard, Imperial Co., Calif., from L. M. Klauber, May 15, 1930. 
Back drab, buffy olive, buffy brown, light drab, or avellaneous. Parotoid same 
color as back. Fore and hind legs same as back, or lighter, such as ecru-drab. 
On sides interstices of green spots are pallid brownish drab, pallid vinaceous- 
drab, or pale drab-gray. Same color on light areas on side of face. Dark dorsal 
spots citrine-drab, deep grape green, or grayish olive outlined by chrysolite 
green edges or deep seafoam green. On sides, warts of green spots have rainette 
green or deep chrysolite green tips and interstices. On dorsum, warts are wood 
brown or avellaneous. Front and rim of upper eyelid vmaceous-fawn. Round 
spot on upper eyelid. Elongate pair inside rear end of parotoid and near 
meson. Another oblique pair in middle of back. Anothersmall pair far back 
and near meson, and another pair far separated just ahead of horizontal of 
vent and just at edge of femur. From rear of parotoid a long spot. From eye 
over tympanum along lower edge of parotoid a spot. This broken but soon 
resumed as long band from shoulder insertion to groin. Between this and 
mesal spots are irregular small spots; so also below this spot. Fore limbs and 
hind limbs with prominent spots, also groin. Dark spot from eye to over 
angle of mouth. Spot below eye. Faint line from eye to nostril to labial edge; 
spot oblique below nostril. Under parts white. Belly and underside of femur 
cream color, warm buff, or colonial buff, becoming pale cinnamon-pink on 
tibia. Throat same color as venter except lower throat under lappet, which is 
vivid brown, vmaceous-drab, or black. Ins rim pale cinnamon-pink or shell 
pink; otherwise black with much spotting of light congo pink. 

Another male. Back drab gray or light cinnamon-drab. All the spots de- 
scribed in other specimen absent or very small. Face uniform. Toes, fingers, 
and membranes somewhat onionskin pink or vinaceous-cmnamon. Other- 
wise like other specimen. 

Female. Back buffy brown. Spotting much as in male. Throat same color as 
venter. No throat apron. 

Structure: Head broad; snout short; sides and front of snout steep; boss 
conspicuous, ridges divaiicating at rear; parotoid glands small, elevated, 
widely separated, oval in shape extending obliquely downward; two meta- 
tarsal tubercles with free cutting dark edge; hind leg approximates body 
length; hind foot long in proportion to hind limb; interorbital space narrow; 
internasal space greater than interorbital; snout only equal to or less than eye; 
toes webbed, webs deeply indented; nostrils set far apart; femur short; horny 



166 HANDBOOK OF FROGS AND TOADS 

excrescences on back of first finger of male, and to less extent or lacking on 
the second finger. 

Voice: The vocal sac of the male is a large "sausage" extending out and up- 
ward. Deflated, the thin discolored skin is closely folded under an apronlike 
extension of light colored pebbly skin of the throat. It is at the rear of the 
throat, and the "apron" may hang down over the forebreast as much as 15 
mm. The call is made up of harsh, low-pitched notes. 

"This form is one of the rarer amphibians of the region. With Bufo alvarms 
it was found almost always in the vicinity of watering troughs. Several were 
caught during the night of July 7 singing on the bank of a pond where the 
large series of Scaphioptis were taken. . . . The song or trill is exceedingly loud 
and very harsh and tinny, reminding one very much of a Klaxon auto horn. 
The interval between songs is rather long, nearly a minute, and as in Bufo 
americanus and Bufo fowleri the trill is sustained for a considerable length of 
time. When singing, the pouch is very pointed and arises under the throat 
and projects forward and upward when fully distended to a point about level 
with the anterior part of ihe lower jaw (A. I. and R. D. Ortenburger, Ariz., 
1926, p. 103). 

Green River, Emery Co., Utah. "Songs of this species and B. woodhousii 
with which it was associated, could be heard in many parts of this little valley 
at night" (V. M. Tanner, Utah, 1928, p. 24). 

Breeding: These toads breed from April to September, dependent upon 
rainfall; in the northern states of their range, from May to July. Little is 
known about the eggs and tadpoles. Amplexation pectoral; egg strings single, 
sometimes double, no inner tubes or partitions seemingly of Bufo compactilis 
type; mature tadpole mottled gray and brown dorsally; light greenish yellow 
and reddish ventrally; tail fin highly arched, obtuse apex; tooth rows %; 
transformation size, 21-28 mm.; tadpole period i%-2% months (largely after 
A. N.Bragg). 

Bragg's studies (1936, 1937? *93Qa) of Bufo cognatns were much needed. 
They mark one of the most significant contributions to anuran life histories, 
in which the Oklahoma group have recently been leading. Bragg gives the 
individual egg diameter as 1.18 mm., the egg inner envelope 0.22 mm. beyond 
the egg, and an outer tube of jelly 2.05 mm. in diameter or 2.66 mm. when 
eggs are in a double row. He gives an egg complement of 20,000, the eggs with 
black animal pole and white vegetative pole. The scalloped outer tube places 
this egg file in the Bufo compactilis class. 

"At hand are two series of tadpoles of this species; one series was collected 
July 2, 1938, 1.5 miles cast of Meade County State Park, Kansas, and the other 
lacks data. The second lot contains numerous sizes of tadpoles from 14 mm. 
to 31 mm., and several transforming specimens which clearly possess the 
pattern so typical of this species. 

"Mouthparts in both series (consisting all told of about 200 specimens) are 



TOADS: BUFONIDAE 167 

fairly constant except in the transforming and extremely young specimens. 
. . . The outer row of teeth of the lower labium is sometimes a little shorter 
or longer than the figure shows, but the average is about as indicated. The 
extent of the medial edge of the papillae on the lower labium varies some- 
what; in some, the papillae barely reach the level of the ends of the outer row 
of teeth, while in others they overlap the ends slightly. 

"Measurements agree with those given by Bragg, except that appearance of 
the hind legs occurs at about 15 mm.; the fore legs appear at about 28 mm. 
A pattern recognizably similar to that of the adult is evident at about 20 mm. 

"These tadpoles show such a striking similarity to those referred by Wright 
to Bufo compactilts Wiegmann . . . that their conspecificity is suggested" 
(H. M. Smith, Kans., 1946, pp. Q5-96). 

Journal notes: July 8, 1917. We found a large stream rushing through Sierra 
Blanca, Tex. They had not had rain for six months. The flat land was over- 
flowed, and a swift current went under the small bridge. At seven o'clock, 
while it was still day, we heard no notes in the creek, but later when we were 
camped one-half mile away, we heard the chorus plainly and decided it must 
be spadcfoots. . . . We found three species of toads and two of spadefoots 
migrating from the mountain sides of Sierra Blanca downward toward the 
pool and noise. The boys, at night, captured B. woodhousn, B. cognatus, and 
B. compactilis on the hillside and in the stream. 

May 20, 1942, Beaver Dam Lodge, Ariz. Big pool north end of bridge. From 
the pool came two very piercing shrill calls. It was B. cognatus. We collected 
the one we heard call and another in the pool In the closed bag its note is a 
harsh nasal clack; that of the tvoodhottsn is a friendly chuckle. 

July 21, El Paso. On road from North Junction toward Smeltertown took 
one or two B. cognatus adults and some three or four young. 

Authorities 9 corner: 

F. A. Hartman, Kans., 1907, p. 228. A. N. Bragg, Okla., 19373, p. 283. 

J. K. Strecker, Jr., Tex., 1910, pp. 19-21. G. A. Moore and C. C. Rigney, Okla., 
H. J. Pack, Utah, 1922, p. 8. 1942, p. 78. 

F. W. King, Ariz., 1932, pp. 175-176. 

Spade-footed Toad, Spadefoot Toad, Sonoran Toad, Western Toad 

Bufo compactilis Wiegmann. Plate XXXIII; Map 8. 

Range: Southern portions of Utah and Nevada, south far into Mexico, and 
east to southwest Oklahoma and the eastern timber belt of Texas. 

Habitat: We found this road breeding in rain pools in open fields near 
streams, in pools in creek valleys, in irrigation tanks or cattle tanks. It is a 
desert form that may at times be seen feeding at night under the street lights 
of desert towns. 

Size: Adults, 2% 2-3% inches. Males, 52-78 mm. Females, 54-91 mm. 



i68 



HANDBOOK OF FROGS AND TOADS 




Plate XXXII. Bufo cognatus (X%)- M- <* XXXIIL B "/ comfaeau. 1,2,5. 

Females. 2,?. Males. Males (X%)- 3- Male croaking (Xw). 4- 

Egg files 



TOADS: BUFONIDAE j6 9 

General appearance: This broad, squatty toad of medium size is pinkish 
drab in color, marked with dull citrine spots. The fingers and toes are light in 
color. The under parts are light. The back is covered with light-tipped tu- 
bercles. 

Color: Beeville, Tex., March 27, 1925. Male. In general color a light drab or 
cinnamon-drab. Tips of the tubercles (under lens) are xanthine orange but 
to naked eye flesh pink, this giving some pinkish tone to the upper parts. 
Spots on back, sides, legs, look dull citrine. Under lens the spots are Prout's 
brown or some other brown with bases of the tubercles mignonette green and 
tips of tubercles xanthine orange. Fingers, toes, and tubercles of fore and hind 
feet pinkish cinnamon. Tip of chin is white, then folded part of throat is 
pinkish buff with ecru-olive in center. This area is followed by a circular pec- 
toral area of purplish lilac. Pupil of eye distinctly horizontal, with horizontal 
black bar through the eye at either end of the pupil. Rim cartridge buff or 
marguerite yellow. Iris spotted vinaceous-buff and bright green-yellow with 
sudan brown vermiculations. Lower eyelid clearly transparent. Tympanum 
dark gray. 

Female. White below. Chin with a few light grayish olive spots. Tubercles, 
fingers, and toes buff-yellow. Back is cinnamon-drab. Upper parts not strik- 
ingly different from male. Tympanum pale olive-gray. 

Structure: Parotoid, elongate, sometimes smooth; no sharp-edged ridge 
from eye to nostril, nostril area smooth; crown without bony ridges, snout 
short, blunt; mterorbital space about equal to upper eyelid; first finger at least 
equal to second; toes half webbed; sole tubercles large, each with a cutting 
edge; tympanum much smaller than the eye. 

Voice: The vocal sac is a large fat oblong "sausage." Deflated, it forms a 
light apron covering several darker folds in the rear of the throat. J. K. 
Strecker, Jr. (Tex., iQ26f, p. 12), calls it "a loud explosive quack." 

May TI, 1925. San Antonio, Tex. At Leon Valley Creek a big chorus of 
B. valliceps, B. debihs, B. compactihs, and S. couchh in considerable num- 
bers. . . . B. compactilis males were calling from the bank. Each one seemed 
to have a favorite perch, and when scared away from it by our light some- 
times would return almost immediately. Their call is loud and shrill and with 
a big chorus would be deafening. 

May 12. No B. compactilis calling at Leon Valley Creek tonight. 

May 29, Comfort, Tex. A few were calling when we first reached the spade- 
foot pond. Their call is very shrill. ... In the pond were many oats half- 
grown or more with oats on the stems. Beside an oat stem where a clump of 
oats might be, a male would rest in water in vertical fashion, sausagelike 
throat extended i%-2 in., directed outward and upward above his head much 
like B. querckus. When a pair is picked up, the male gives a call like a pea 
fowl or hen, clucking much of the time. The water was very shallow where 
I caught the pair. 



170 HANDBOOK OF FROGS AND TOADS 

June 15. Left Rio Grande City 7:45 A.M. Stopped at Santa Cruz ranch op- 
posite hill with U.S. Geological Survey marker. At top of hill we could clearly 
hear B. compactilis in a pond across the road. These males which are croaking 
in the daytime are in the dense shade of some overhanging trees. 

Breeding: These toads breed from May i to July 10, with a few stragglers 
later at the time of the late summer rains. The brown and yellow eggs are in 
long fine coils; the jelly tube narrow, %2 inch (2 mm.); the eggs crowded, 
14-20 eggs in i% inches (30 mm.); and the vitelli Ko inch (1.4 mm.). The 
eggs hatch in two days. The bicolored tadpole is small, 1-1% inches (24-28 
mm.), light-colored, its back a drab or light grayish olive and its belly pale 
cinnamon pink; its tail crests are translucent. The tooth ridges are %. After 
40 to 60 days, the tadpoles transform, June i to August i, at % inch (12 mm.). 

Journal notes: March 22, 1925. Started for Corpus Christi, Tex., at 6 P.M. 
Captured B. compactilis, north of Kansas City. A female crossing the road. 

March 25. Tonight took two Bufo compactilis males. In no place are they 
breeding so far as we can see. Tonight we picked up what we at first took for 
a toad of B. compactilis build. Not until later did we realize we had a spade- 
foot. 

March 26. Tonight in Beeville, Tex., under electric light saw a female mov- 
ing along. Caught her. 

May 29. Comfort had 3 inches of rain in the afternoon of May 28. We went 
to San Antonio and at 5:30 P.M. started for Comfort. At Big Joshua Creek 
evidences of flood but no frogs calling. . . . Left Boerne and arrived at Com- 
fort about 8:30 P.M. The Guadalupe River had been up high but was down 
at this time. No frogs at all. When we pulled up to Comfort filling station, 
heard a great chorus south of the filling station. We went down from the main 
road south toward baseball field, then left toward the creek. It was slippery. 
We crossed creek and in a cultivated field ^-% mile farther on was a great 
volume of "wows or meows 1 ' or catcalling (S. cottchit). In this pond were 
many B. compacttlis. A few were calling near the edge but more in the water. 
... At the far end of the cultivated field the pond extended into a grassy 
pasture. Here the B. compactilis were calling in considerable numbers. A few 
had standings on the shore, several were out in clumps of grass. They are well 
called the spadefoot toad. They are usually the associate of the spadefoot (S. 
couchit) . 

We left this pool and went on to Cypress Creek where there are water lilies. 
Nothing doing. We went on 2 or 3 miles, then returned. Down a side road 
heard S. couchii. Found pools beside the road. Herein were many S. couchti 
and many B. compactilis. 

June 7. As we approached Encinal, close to the road in two pools were many 
small tads toad tads, but not spadefoots. We had lost our net but found 
strainers good for catching them. Are the toads B. compactilis or B. debtlis? 
When the pools begin to dry up, gracklcs go there to eat up tads. 



TOADS: BUFONIDAE 



171 



June 9. At Dolores, Tex., collected many Couch's spadefoots and B. com- 
pactilis in large concrete-bottomed irrigation tank. Around the edges are a 
few places where mud has gathered, and in such places B. compacttlis has dug 
into the mud sometimes leaving an opening, sometimes just leaving the 
earth rough. 

June 2, 1942. Beyond Rockville in Virgin River saw countless small Bufo 
tadpoles. Two to three miles from Springdale, Utah, I went after toads in the 
dark. Had to jump a drainage ditch. In a cultivated field no end of water; one 
pool. I thought I had Scaphiopus but the call was not so harsh but much 
harsher than the few Bufo woodhousii then calling. Caught five female 
B. compacttlis. When I approached, the males stopped; never saw one. We 
saw a Bufo compactihs crossing the road in Rockville. 

July 22, 1942. Toyahvale, Tex. Night trip west on highway. Heard one 
Bufo. This proved to be Bufo compactihs. 

Authorities' corner: 

J. K. Strecker, Jr., Tex., igoSb, p. 82. J. K. Strecker, Jr., Tex., 1926^ p. 12. 

J. K. Strecker, Jr., Tex., 1915, p. 52. J. K. Strecker, Jr., Tex., 1928^ p. 7. 

G. P. Englehardt, Utah, 1918, pp. 78- H. M. Smith, Gen., 1947, pp. 7-13. 

79- 

In 1940 Linsdale (Nev.) wrote: "It is necessary, in making final taxonomic 
appraisal of toads, to study the living animals in their natural surroundings. 
Until further field studies can be made on this group of forms, satisfactory 
conclusions cannot be reached. However, the preserved material is sufficient 
to show that some changes in systematic treatment of the group will be re- 
quired. Present indications are that these toads are all in the same species. The 
oldest name in the lot is compactihs. At least four races seem to deserve recog- 
nition, as follows: Bufo compactihs compactilis Wiegmann 1833, Bufo com- 
pacttlis woodhousii Girard 1856, Bufo compacttlis fowlen Hmckley 1882, 
Bufo compactihs cahfornicus Camp 1915." 

He may not be far from the right solution but we cannot yet accept this ap- 
praisal without more study of the western forms in the field. Should Bufo 
cahfornicus be placed as a Bufo compactihs subspecies rather than as a Bufo 
cognatus affiliate? Is Linsdale's Bufo compactihs of Nevada the same as the 
large Bufo compactihs of Texas? In 1942 Dr. Klauber remarked that he 
wished we could solve the B. compactihs problem. The Texas Bufo com- 
pactilis is an old acquaintance, but the western representatives we do not 
know well in the field. 

In a recent paper H. H. Smith (Gen., 1947) divides B. compacttlis into two 
forms restricting B. c. compactihs to the main Mexican plateau and B. c. spe- 
ciosus for the U.S. form. We are glad he did not set off the Nevada and 
Arizona form as new. The discontinuous range of his B. c. speciosus may not 
be real. Field work in the interval in Arizona may yet produce this burrower. 



172 



HANDBOOK OF FROGS AND TOADS 



Little Green Toad, Sonoran Toad, Green Toad, Sonora Toad 

Bufo debtlis Girard. Plate XXXIV; Map 12. 

Range: "Eastern Texas and Tamaulipas" (Taylor, 1936). "In the lower 
part of the valley of the Rio Bravo (Rio Grande del Norte) and in the prov- 
ince of Tamauhpas" (Girard, 1854; Stejneger and Barbour, 1938). 

Habitat: On grassy mesquite flats, breeding in temporary rain pools, 
ditches, or shallow pools in streams of intermittent flow. 

Size: Adults, i-i 4 /f, inches. Males, 26-41 mm. Females, 31.5-46 mm. 




Map 12 

General appearance: This beautiful little green toad has black spots on the 
back and some gold- or yellow-tipped tubercles on the sides and legs. The 
legs have a few dark barhke spots with light central areas. The legs are short, 
the snout pointed, the parotoids large and elongate or triangular, the body 
narrow and rounded in contour, the head narrow with interorbital space pro- 4 
portionally wide. There is a black bar at the arm insertion and there may be 
a black line or two or three spots in the middle of the lower breast. 

Color: Male. Alice, Tex., March 31, 1925. Back spinach or cedar green or 
hellebore green to calliste green or cosse green. Back with black spots and 
black tubercles on these spots. On the sides, besides green and black, some 
tubercles are light cadmium, lemon chrome, or maize yellow. These colors 
also appear on dorsum of hind legs and brachium of fore limb, and on web- 
bing of hind feet and tubercles. The rear of thigh is green, black, and light 
cadmium color. There is a line of light cadmium where dark colors meet the 



TOADS: BUFONIDAE 



'73 



white under parts. Some of the tubercles on greenish background with light 
cadmium tips. Light buff or cream-buff along upper jaw to arm insertion. 
Rim of lower jaw has a white margin followed by dense black, then pale olive- 
gray to deep olive-gray. There is a black bar on the venter in the pectoral 
region opposite arm insertion; also black on the brachial insertion. Sometimes 
there is a black line in the middle of lower breast or two or three black spots. 
Iris is mainly black with some pale ochraceous-buff. 

Female. June 15, 1925. In general the same as male, but without the black 
chin. Under parts are olive-buff or white. In some the two pectoral black 
spots are absent or not so well developed as in the male. 

Structure: Crown without bony ridges; snout protruding; first finger 
shorter than the second; hind limbs usually shorter than body length; tym- 
panum small; toes slightly webbed at base; foot short; no tarsal fold; paro- 
toids sometimes extending downward to the level of the jaw. 

In view of Taylor's (1936) concept, the following Strecker quotation is 
composite, i.e., for both species: "Structure: Girard's original description 
(May, 1854) of Bttfo debilts (p. 87) varies so slightly from his Bitfo insidior 
(p. 88) that we include it for future clarification of the status of each form. 
The description is: Upper surface of head without any crest on ridge. Snout 
rounded. Mouth moderate. Upper jaw emarginated. Tongue small. Tympa- 
num small. Parotoids moderate and elongated. Limbs of moderate develop- 
ment, femur shorter than tibia. First finger longer than the second. A large 
metacarpal disk. Toes slightly united at the base by a web. Two metatarsal 
processes. No membranous fold at the inner lower margin of the tarsus. Skin 
above pustulous; pustules of moderate development; warts beneath very 
small. Color above brownish yellow, spotted. No dorsal lighter vitta. Beneath 
of a uniform soiled yellow. Allied to Bujo speciosits. . . ." 

Voice: The vocal pouch is a round ball reaching to the tip of the chin. The 
call is cricketlike, a low sustained trill. 

March 28, 1925. Between San Diego and Alice, Tex., we heard a note abso- 
lutely new to us something like a cricket. When we came to the pond and 
it appeared light-colored under the flashlight, I knew it must be Btifo debilis. 
It is rightly named "the little green toad." In some ways its note makes me 
think of Fowler's toad, but it is not nearly so loud or strong. The call is sus- 
tained. When I first went down to the pond, I thought I heard only one, but 
before I got the light on it to see how it croaked, two of them started hopping 
toward the ditch. In some ways, the toad and its note remind me of B. qtierci- 
cus f but B. quercicus is much louder and shriller. 

When held in hand the male gave a little batlike click, like marbles hitting 
together. Over in the mesquite, "devil's elbow," and prickly pear, we went 
for two or three we heard, and came back with one of them. There was a small 
ditch or runway there. 

In a shallow overflowed mesquite area, several B. debilis males were calling. 



174 HANDBOOK OF FROGS AND TOADS 

Oftentimes when you approach, or particularly when you put light on them, 
they duck flat to the ground. Bufo debilis note at times something like the 
"clucking" one gives to make a horse travel faster. 

Breeding: This species breeds from the last of March to mid-June. "The 
eggs are in small strings and are attached to grass and weed stems. . . . The 
tadpoles are slightly smaller than those of B. punctatus" (J. K. Strecker, Jr., 
Tex., 1926^ p. 10). They transform at %-% inch (8-11 mm.). 

Couch at Matamoras, Mexico, took six specimens (USNM no. 2621) which 
measured respectively 8.0, 8.5, 9.0, 10.5, ii.o, n.o mm. This probably repre- 
sents a close approach to transformation. Doubtless this species transforms at 
a little larger size than B. quercictis, of which the smallest transformed indi- 
vidual recorded is 7 mm. 

"Their metamorphosis is accomplished within a very short space of time. 
This is necessary on account of the extremely temporary character of their 
breeding places. I returned twenty days later to one pond in which I had 
found debilis breeding and discovered that it was almost perfectly dry, only 
a few mud holes remaining to indicate moisture. In one of these mud holes 
were a few belated tadpoles, and in the grass along the banks I found two 
small toads with tails" (J. K. Strecker, Jr., Tex., 1926^ pp. 10-11). 

Journal notes: April 3, 1925, Gardendale, Tex. In a shallow pool near the 
station were some small Bufo tadpoles. Returned tonight at 8 130 P.M. Air at 
least 72. Heard 3 or 4 Bufo debtlis. 

May 9, San Antonio, Tex. At Leon Creek, several Microhyla, and one Bufo 
debilis. This one we photographed. He did not duck under the flashlight. In 
the next valley bottom in gravel pit stream several B. valliceps, one Micro- 
hyla, and one B. debilis. 

May u, San Antonio, Tex. Went to Somerset in the afternoon and "spot- 
ted" ponds by the roadside. Back to the city for supper; then out Somerset 
road again. First stop on return 7:45 at roadside pond now swelled by rain. 
Heard lots of Pseudacns, meadow frogs, Hyla versicolor, Bufo valliceps. Bufo 
valliceps are calling from temporary islands and the banks. Little nearer the 
city, beside road nothing but Microhyla. Next stop roadside pond 25 male 
Bufo debilis ', sometimes ^ or 4 together at edge of plowed ground 3 or 4 Man- 
cy's garters evidently enemies of all of them. Great chorus of Microhyla and 
Pseudacris. 

Jan. 18, 1942. Visited Al Kirn of Somerset. Showed me his material plenty 
of Bufo debilis. These he loaned me. 

Authorities' corner: 

J. K. Strecker, Jr., Tex., ipoSb, p. 81. E. H. Taylor, Kans., 1929, p. 445. 
J. K. Strecker, Jr., Tex., 1915, p. 51. E. H. Taylor, Gen., 19363, p. 513. 

R. Kellogg, Gen., 1932, p. 50. 



TOADS: BUFONIDAE 



175 




Plate XXXIV. Bufo debilis. i. Female 
(XO. 2. Male croaking (X%)- 3- M*'e 
(X%). 4,5- Females (X%)- 6. Female 



Plate XXXV. Bufo exsul (X%). 
Young. 2,7,8. Females. 3,4,5,6. Males. 



176 HANDBOOK OF FROGS AND TOADS 

Black Toad 
Bufo exsul Myers. Plate XXXV; Map n. 

Range and Habitat: Known only from Deep Springs. "Deep Springs Val- 
ley is an isolated depression in the desert mountains of northeastern Inyo 
County, California. It is elongate in form, trending northeast by southwest, 
about twelve miles long and five miles broad at its widest part. The lowest 
part of the valley, at its wide southwestern end, is a flat area of about three by 
five miles, of almost exactly 5,000 feet elevation, although the rest of the gently 
rising valley floor is also very level. Surrounding Deep Springs, the White and 
Inyo Mountains rise to heights of 7,000 to 8,000 feet. Westgard Pass, through 
which one enters the valley from Owens Valley to the west, reaches 7,276 feet, 
and the top of the pass into the southern arm of Fish Lake Valley, on the east, 
is* at 6,374 ^ eet - The lowest entrance to the valley appears to be the dry, narrow, 
and now virtually unused Soldier Pass, from the dry northeastern corner of 
Deep Springs Valley into Eureka Valley on the southeast; the top of this pass 
appears to be at approximately 5,400 feet. 

"Like other desert valleys to the east of the Sierra Nevada, Deep Springs 
is exceedingly dry, and on its floor the vegetation consists of sparse low desert 
brush (Chrysothamnns}. The surrounding mountains support growths of 
juniper and pinon. 

"The valley has few sources of water. Aside from washes carrying water 
only during infrequent desert rains, I know of only three. Wyman Creek, the 
course of which leads into the northern end of the valley, contains a little wa- 
ter, at least in its upper reaches, most of the year, and tiny Antelope Spring, 
on the west side of the valley, appears to be permanent, but neither of these 
contributes water to the valley floor, except during exceptionally heavy rains. 
The chief water source is formed by the Buckhorn or Deep Springs which 
flow from the base of the southeastern valley wall just above the sink. These 
springs issue from the rocks for a distance of a mile or more, but only a few 
of them have a strong flow. The flow from the more southerly springs forms a 
marshy area of several acres, the water finally draining down into a shallow 
lake of alkaline, sulphurous water that sometimes reaches a diameter of a mik 
or more. There is also a smaller pond of good water between the springs and 
the lake. The marshy area and the watercourses through it emit a strong sul- 
phurous odor. Much of this is apparently due to sulphur bacteria, but some of 
the springs themselves must carry sulphur" (G. S. Myers, Calif., 1942, pp. 

1-2). 

Size: Adults i%-2% inches. Males 44-59 mm., average 52 mm., mean 51 
mm.; females 46-61.5 mm., average 54 mm. (Measurements of 51 specimens 
of the Ferris- Wiggans-Myers accession May i, 1937, and of the C. L. Hubbs 
family accession Sept. 4, 1934.) 

Of 25 adults in our collection from Deep Springs Valley School, May 12, 



TOADS: BUFONIDAE 



177 



1942, we have the following sizes: adults 45-56 mm.; males (17) 45-53.5 mm., 
average 49 mm., mean 48 mm.; females (8) 45(?)~56 mm., average 49 mm., 
mean 51 mm. There were five questionable as to sex, 39-43.5 mm. The inter- 
mediates measured 29-40 mm., except for one 25.5 mm., average 33 mm., 
mean 34 mm. What is the transformation size? 

General appearance: "A localized derivative of Bttfo boreas, closely similar 
to B. b. nelsoni in its small size, narrow head, and smoothness of skin, but 
sharply distinguished from nelsoni, from all forms of boreas, and, indeed, 
from any other North American Btifo by its strange color. The dark dorsal 
markings of boreas have enlarged, fused, and darkened until the upper sur- 
faces are almost entirely a shining lacquer black (deep dull blackish brown 
in alcohol; grayish black in formalin), the remnants of the light interspaces 
remaining as irregular, whitish, or brownish vermiform markings, and the 
vertebral line showing as a white or whitish hairline, frequently greatly inter- 
rupted, down the middle of the back. The markings of the underside are even 
more remarkable and diagnostic. The sparse black spots of boreas boreas or 
boreas nelsoni have developed into a dense mottling or marbling of black, 
which appears in life as if made with india ink. Not only are the belly and 
lower surfaces of the tibiae and tarso-metatarsals heavily marbled, but the 
throat is usually spotted, and the undersides of the femurs and rump are com- 
pletely black except for the tubercles, which are white. The tarsal fold is very 
poorly developed and is almost obsolete in most of the specimens" (G. S. 
Myers, Calif., 1942, p. 3). 

Color: Deep Springs, Calif., May 12, 1942. Young, half grown. Middorsal 
stripe is primrose yellow m one and marguerite yellow in another. The paro- 
toid in one bears a prominent patch of isabella color, another of olive ocher. 
This color extends down the back between tubercles and becomes lighter, 
sometimes deep seafoam green. The general background of back is light 
brownish olive with about three irregular rows of warts, one on either side of 
middorsum, one back from parotoid, and an intermediate one. Bordering 
this light brownish olive area is a broken irregular thread of marguerite yel- 
low, below which is a band of almost clear black along the sides. The black 
of the lower sides is broken into mottling by marguerite yellow. Marguerite 
yellow or ivory yellow is the dominant ventral color, very heavily spotted with 
black so heavily spotted on rear of belly and across thighs that the nickname 
"black pants" was used in the laboratory. On the face is an oblique band of 
ivory yellow or marguerite yellow extending backward from the eye, some- 
times forming a half moon and reaching the angle of the mouth. There is a 
black band from eye to arm insertion, bordered behind by a longitudinal 
stripe of marguerite yellow or a broken row of spots of the same. The iris is 
black with prominent speckings of glass green and with a wash of cinnamon 
in the upper portion; the pupil rim is of reed yellow. The young are very sug- 
gestive of the females of B. canorus. 



178 HANDBOOK OF FROGS AND TOADS 

Female. The female has very low tubercles, smooth and flattened, the mid- 
dorsal stripe of the young, and a few flecks of marguerite yellow along the 
sides (otherwise the dorsum is entirely black). These toads have the most 
prominent marguerite yellow and black markings on the upper surfaces of 
fore and hind limbs, particularly on the underside of tibia and dorsum of foot. 
The underside of the foot is black, except for the articular tubercles which 
are light yellowish olive. The throat is more heavily spotted than in the young. 
Iris as in the young. The face marking has the oblique light line, then black 
area from eye down, then irregular light longitudinal stripe back of angle of 
mouth. 

Male. No striking difference from female. The inner palmar tubercle is very 
prominent. 

This lack of sexual dimorphism makes these toads different from B. cano- 
rus, but in general appearance they seem like a smaller black B. canorus. 

Some random notes on the Ferris, Wiggans, and Myers material follow: 
All except one male have thin light stripe full length of body. One female has 
line absent in rear half and broken in the front half. One female has line 
broken in many places. The young have a prominent stripe. The first young 
to have uniform black back is 40 mm. long (no. 2553) ; those 36 mm. and 38 
mm. are spotted on either side of middorsum. Even some of smallest have 
black on sides. One of striking things in this form is the thin light line going 
through interorbital area and the oblique bar crossing from one eyelid to the 
other. In many this cross thread is absent and there is a light longitudinal spot 
on parotoid, but this is absent in some. The venter is heavily mottled with 
black from pectoral to pelvic region; the groin and underside of femur and 
pelvic region are black with white pebbling. On forward half of venter the 
white and black are about equally divided. The throats are lighter, are creamy, 
but with black clots in most cases. Six of the small ones have scant spotting on 
throat region and four with heavy dotting. All ten of these have prominent 
black pectoral spot. Some of females look ripe. 

Structure: "Description of Holotype. Head rather narrow, its width approx- 
imately three times in length to vent, with a weak canthal angle. Snout moder- 
ately rounded when viewed from above, the eyes projecting somewhat beyond, 
the line of the upper lip. In profile the snout is bluntly rounded but slop- 
ing and not vertically truncated, the upper lip being the most anterior point. 
Nostrils slightly nearer to eye than to tip of upper lip, the distance from the 
eye equal to internarial space and to interocular space, which is flat. No cranial 
crests. Loreal region somewhat inclined, slightly concave. Depth of subocular 
area equals half length of exposed part of eye. Distance from nares to tip of 
upper lip equals length of exposed part of eye. Tympanum indistinct, its 
upper posterior border partly obliterated, vertically oval in form, its depth 
equal to distance from nares to eye, or not quite two-thirds horizontal diame- 
ter of orbit. Tympanum close to eye, its distance from latter scarcely equal to 



TOADS: BUFONIDAE 179 

the narrower (horizontal) diameter of tympanum. Distance of tympanum 
from the corner of the mouth (directly below it) equals slightly more than 
.(vertical) depth of tympanum. Parotoid glands moderate, oval to triangular, 
wider and more distinct posteriorly than anteriorly, where they fade out just 
before reaching eyes, their length approximately equal to distance from eye 
to tip of upper lip (snout tip) ; the glands are much farther apart than their 
own width and do not descend to the sides appreciably behind the tympanum. 

"Arms moderately stout, the fingers entirely free of web. Third finger long- 
est; second and fourth approximately equal, reaching base of penultimate 
phalanx of third. First finger equal to or barely surpassing second. Subarticu- 
lar tubercles single, well developed only at base of each finger. Several small, 
scattered tubercles on each palm. Two distinct, enlarged palmar tubercles, the 
outer large and rounded, its highest point toward its distal end, the inner 
about half the size of the outer and somewhat more convex. 

"Legs stout and relatively short, the tarso-metataisal joint reaching middle 
of tympanum when leg is brought forward, the heels not quite touching when 
femur and folded tibia are brought to right angles with body. Toes more than 
half webbed, the web reaching as far as the base of the antepenultimate pha- 
lanx of the fourth toe and the penultimate phalanges of the other toes; the webs 
are not greatly excised. Inner metatarsal tubercle moderate, elongate oval in 
form, its end only slightly free and without cutting edge; the tarsal fold run- 
ning proximally from the tubercle is nearly obsolete and only barely visible. 
Outer metatarsal tubercle smaller, rather conical and rounded. Subarticular 
tubercles of toes single but weak" (G. S. Myers, Calif., 1942, pp. 4-5). 

Comparison of comparable Bufo canonts and Bttfo exsul t each at 44 and 56 
mm., shows head wider, eye and interorbital space slightly larger, tympanum 
much larger, fore limb longer in B. canoms, and third toe slightly longer in 
B. exsul. One must remember that the Bufo canorns is hardly mature at 44 
mm. and reaches 75 mm. in extreme size, whereas B. exsul is mature at 44 and 
reaches only 61.5 mm. in extreme size. 

Breeding: On May 12, 1942, two different sizes of tadpoles were with the 
adults: many tadpoles were 19 or 20 mm. in length, and a mature tadpole set 
were 29-35 mm., with body lengths such as 12.5, 13.0, 13.5, 13.5, 14.0, and 15.0 
mm. for six we measured. The tooth formulae are %. The tail tip is very 
rounded. 

Journal notes: May 12, 1942. Went to Deep Springs. At 12:30 arrived at 
Springs, 5000 feet elevation 7 Mi miles south of- Deep Springs School and above 
Deep Springs Lake. Just below the first spring, in the water runways between 
tussocks, began to see Bufo exsul. The bottoms are mud, water 2 to 4 inches 
deep. These areas more or less shaded by tussocks. Soil dark and mucky. De- 
pressions between tussocks 8 inches to i ! / feet deep. The toads often dashed 
into holes in tussocks or into shaded runways. The first three running springs 
unite into one clear stream. In the stony clear sources no toads; just above 



180 HANDBOOK OF FROGS AND TOADS 

confluence of three springs, where streams are not so fast, among some rushes 
and muck, several of the biggest ones were found. One ran into a hole under 
a tussock. Along this stream 4 feet wide and nowhere more than 2 feet deep, 
usually i foot or less, were many toads. Every so often toads would swim out 
from bank and stay poised on top. Often they would swim downward and 
hide in vegetation. Occasionally they would rush into the mud but seldom 
completely out of sight. Sometimes one could see in one area as many as 3 or 
4 toads on bottom or poised in water below surface. Often a shaded spot or 
place with no shade would have 2, 3, 4, or even 6 toads in one localized area. 
Are they approaching breeding? No mated pairs. In this area one also could 
see toads in the vegetation at surface or see one or two poised on a mat of algae 
at the surface. The farthest from the stream we took toads was 6 feet. Even 
near stream they made for stream and escaped in it. 

In the stream were three different sizes of tadpoles. Some seemed mature 
but no transformation sizes were available. The warm sulphur spring beyond 
had just as many toads. Some of the largest ones are blacker, some of the 
smaller ones were somewhat yellowish. Mr. Dieflfenderfer said he and another 
found 2800 of them frozen in the water of Antelope Spring west of Deep 
Springs Valley Lake. Did the big tadpoles come from breeding in March this 
year? 

May 13. Started about 8:30 from Deep Springs. Snow all about. 

Dr. Myers showed us his and the Hubbs material from Deep Springs pre- 
vious to the publication of his new form (September, 1942). 

Authorities 9 corner: 
G. S. Myers, Calif., 1942, pp. 8-9. 

Canadian Toad, Winnipeg Toad, Manitoba Toad 

Bttfo hemiophrys Cope. Plate XXXVI; Map 10. 

Range: North Dakota, Manitoba, Alberta to northwest provinces. In coun- 
try tributary to Lake Winnipeg in Canada. Some range discussions include 
Minnesota, South Dakota, Montana, and Mackenzie. 

Habitat: Stream valleys, lakes, and Turtle Mountain region. 

Size: Adults, 2^4-3% inches. Males, 56-68 mm. Females, 56-80 mm. 

General appearance: These are brownish or greenish toads of medium size 
and with short snouts. The most marked characteristic is the boss on the head. 
This is a narrow raised horny structure extending from the rear of the upper 
eyelids to the snout. Frequently the rear end of this boss forms quite a promi- 
nence. It makes one think of a frontal plate of the upper mandible of some 
rail and cootlike birds. Their backs are covered with many fine, rounded 
warts, generally two to five to a dark area. There are many spiny warts on 
the femur. There is a light stripe down the middle of the back and a conspicu- 
ous dark band along the side, bordered above by a light line. The under parts 



TOADS: BUFONIDAE 181 

are cream or buflfy with numerous small black spots, a prominent one in the 
middle of lower throat. The throat of the male is dark. The broad upper eye- 
lids with closely set tubercles are in sharp contrast with the narrow inter- 
orbital. This makes the eyes stand out. 

Color: Walhalla, N.D., Sept. i, 1930. Male. The stripe down the middle of 
the back is white or ivory yellow, edged on either side with reed yellow. From 
eye backward over lower edge of parotoid and along side to groin extends a 
cream-buff line, becoming chamois on lower edge of parotoid and in vertical 
bar that extends down to the arm. Below the dark lateral band, the interstices 
among the dark spots are the same color. The color of the back in its lighter 
portions is ecru-olive; in darker portions, it is yellowish olive or dark greenish 
olive. The dark bar on either upper eyelid, the area just inside the inner tip 
of parotoid, and the succeeding paired spots along median line are dark green- 
ish olive or olive with brownish olive warts. The band on side of body is also 
dark greenish olive or dark olive. The dark oblique bar, below eye, and big 
bar on shoulder insertion are the same color. The tympanum and interstices 
between these bars are malachite green. The arms and legs arc barred with the 
dark color of the dorsal spots, with interstices the color of back and cream 
color. The rear of femur has the dorsal background color with strontian yel- 
low or wax yellow spots. This color comes into the color of the lower belly and 
underside of femur. The belly is cream-buff, heavily marked with light gray- 
ish olive. In middle of pectoral region and on lower throat is long bar spot of 
the same. The throat is suffused with light grayish olive. Upper portion of 
parotoid is drab. The boss is argus brown. The inner metatarsal tubercle is 
mummy brown. The upper surfaces of first two fingers are covered with ex- 
crescences of same color or of chestnut-brown. The eye is olivaceous black or 
dark ivy green with prominent lemon yellow ins ring; upper part of iris is 
heavily spotted with same color, lower slightly spotted, and upper and lower 
parts slightly spotted with russet-vmaceous. 

Female, young. The spots on belly are larger; the throat is clear of spots ex- 
cept for pectoral spot. Throat is cream-colored. First two fingers without ex- 
crescences. Otherwise marked as in the male. The outer circle of the iris bears 
a deep bluish glaucous ring. 

(From preserved material in USNM.) Back brown with a median light 
band. A broad brown lateral band starts back of the arm insertion extending 
backward to the groin and forward as a broken band to the eye, above the 
tympanum. This is bordered above by a light margin, which sends a series of 
more or less vertical lines to the arm insertion, the mam branch going forward 
involving the parotoid. This dark band borders the light color of the belly, 
which is strongly mottled with dark along the sides. The median line is bor- 
dered on either side by a dark area more or less continuous to the vent. In 
some of the older frogs this breaks up into detached spots about some of the 
collections of warts. The warts are lighter colored. Between the dorsal series 



182 HANDBOOK OK FROGS AND TOADS 

of brown spots and the lateral band are two rows of spots, the more regular 
one being above the light margin of the dark lateral band. There is a brown 
area back of the eye and below the tympanum, a brown suborbital spot, and 
a bar from nostril downward; between this bar and suborbital spot is another 
bar. Therefore the upper lip is an alternation of light snout, nasal bar, light 
interval, dark bar, light interval, suborbital dark spot, light interval, and sub- 
tympanic spot. There is a light line from below the tympanum or near angle 
of the mouth, backward over the arm insertion; it, with the vertical bar from 
the light side band, extends to the dorsum of the arm. The upper eyelid has 
a prominent brown transverse band, margined in front and below by a con- 
spicuous light gray margin. This eyelid coloration stands out in sharp contrast 
to the boss of the head. Breast and belly spotted with dark. The hind legs have 
dark cross bands with reddish tubercles in the center of cross bands. Dark 
reticulations prominent on back of femur. 

Structure: Boss on head from snout to rear of interorbital space; slight fur- 
row down the middle and outer ridges forming boss are parallel; these ridges 
connected by a bar at rear and with slight extension toward the eye; broad 
upper eyelid with closely set tubercles, in sharp contrast with narrow inter- 
orbital; snout short, vertical in profile; tympanum elliptical and vertical; two 
cutting metatarsal tubercles, inner large and spadelike; ventro-basal portion 
of femur discolored; fore throat of male discolored. 

Voice: "Its spring note is a soft trilling, uttered about twice a minute and 
lasting about three seconds" (E. T. Seton, Man., 1918, p. 83). 

Breeding: This species breeds from May onward, in shallow edges of lakes 
or ponds or other water. There are no descriptions of the tadpole on record. 
They transform at %-"!/> inch (9.0-13.5 mm.). 

Journal notes: August 30, 19 30. At n o'clock we started northward from 
Grand Forks. At Ardock we found a stream dried up except for isolated 
pools. Around one in the cow-punched and cracked mud, we took some seven 
or eight young of Btijo hemiophrys. They were of two sets: 36 mm., 33 mm., 
26 mm., 23 mm., 22 mm., and 18 mm. All of these young specimens have a 
suggestion of the dark pectoral spot, and only the two larger ones begin to 
show cranial ridges. 

August 31, 1930. It rained in the afternoon. We went through Neche to 
Walhalla, N.D., at the edge of Pembina Mountains. We arrived about 5:30 
P.M. It was still raining. We began to get desperate in our desire for live adults 
of Biifo hemiophrys. Before dark we took an excursion around town looking 
for possible covers for toads. The evening was cold, the rain had stopped, and 
it was doubtful if toads were abroad feeding. This has been a summer of re- 
markable drought. We used one of the best expedients of a collector. We en- 
gaged a bright-eyed young boy (Audrey Miller) to search the town. Inside 
of half an hour he appeared with an adult male. He and another lad started 
promptly again with a flashlight and returned twice, each time with an adult 



TOADS: BUFONIDAE 



183 








Plate XXXVI. Bufo hcmiophrys. i. Male Plate XXXVll Bufo msid.or (X%) 

(X%). 2,4,5. Males (X%)- 3- Young Males, from Toyahvale, Tex. 

/i .n / \ 



184 HANDBOOK OF FROGS AND TOADS 

toad. We asked where they got them and he said in the gutters, and later he 
said he got them in the sewers. 

Sept. i, 1930. We went out with the boys to a place where they had a toad 
located, but had missed him last night. We wondered why they carried a flash- 
light and a pail on a long rope. We soon found out. The town has drainage 
manholes. The boys took off the cover and with flashlight searched the surface 
of the water and the walls of the hole. They pointed out a toad floating at sur- 
face. They let down their pail and drew it up under the toad and so caught it. 
They brought up two toads instead of one, but one was dead. Quite likely the 
toads were caught here as in a trap, but at this long period of drought it was 
our best chance of getting material. 

These males have a throat of the Bufo americantts class, but close examina- 
tion of one which half inflated its throat revealed that the lower throat region 
expanded first, and there also seemed to be a secondary swelling at the ventral 
base of each arm. The throat seems to have a slight fold across the lower throat 
on the pectoral region. Its boss, this slight indication of a lower throat pouch, 
and two cutting mctatarsal tubercles suggest relationship with B. cognattts. 

Authorities' corner: 
E. T, Seton, Man., 1918, p. 83. 

Sonoran Toad 

Bufo tnsidior Girard. Plate XXXVII; Map 12. 

Range: Colorado and Kansas south through New Mexico, Arizona, and 
Texas into northern Mexico. 

Habitat and Size: Since Dr. Taylor's (1936) resurrection of this Girard spe- 
cies from the synonymy of Bufo debihs and since the inclusion of it in the 
check lists (Stejnegcr and Harbour), little material has appeared to add to 
our understanding of it. 

General appearance: From our limited experience with this form in Trans- 
Pecos, Tex., we would call it more bleached than are southern Texan little 
green toads. 

"The absence of cranial crests, or the presence of very narrow ones closely 
approximated to orbits, coupled with the broad, elongate parotoid gland and 
greenish coloration is very distinctive of this Kansas toad" (H. M. Smith, 
Kans., 1934, p. 443). 

Color: July 22, 1942, Toyahvale, Tex. This toad has a flat head, pointed 
snout, and very large, elongate, and broad parotoids. The back of one is 
courge green, with the top of the head clear yellow-green. Another has a bis- 
cay green back with top of snout and upper eyelids touched with old gold. 
The back and limbs bear tubercles encircled with mustard yellow and tipped 
with mummy brown. The back and top of head are also flecked with black, 



TOADS: BUFONIDAE 185 

each spot with its black central tubercle. Some spots are dumbbells in shape. 
A prominent oblique black bar crosses the eyelid. The hind limbs have two 
or three bars on femur, two on tibia, and one on metatarsus. On the sides the 
tubercles, encircled with chartreuse yellow and tipped with black or mummy 
brown, are on a mummy brown background, which forms a broken line across 
the rear of parotoid and along the side. The groin, underside of thighs, and 
sides of belly are pale brownish vinaceous; the center of the belly is light glau- 
cous-blue, one immaculate, another with several spots. The throat of the fe- 
male is white, that of the male discolored, the forward part deep purplish 
vinaceous, the rear a pale vinaceous-lilac, the middle deep medici blue. The 
underside of hand and of toes is vinaceous-cinnamon, toes and fingers tipped 
with cinnamon-brown. The two metatarsal tubercles and the palmar tubercle 
are russet. The black eye has a citron yellow pupil rim broken in four places, 
and the iris is heavily flecked with ochraccous-buff. 

Structure: The original description (C. Girard, Gen., 1856, p. 88) is as fol- 
lows: "Upper surface of head plane and smooth. Snout subacute, protruding. 
Mouth moderate, upper jaw slightly emarginated. Tongue elongated tapering 
towards both ends. Tympanum inconspicuous. Parotoids large and elongated, 
situated obliquely upon the shoulder. Limbs moderate. First finger equal to 
the second in length. A metacarpal disk, and a tubercle. Toes slightly webbed 
at base. Two metatarsal tubercles. Skin papillous above, warty beneath." 

Voice: "This species is rather abundant throughout the region traversed 
(eastern border of the Staked Plain of Texas) . It is frequently found in the 
grass, where ils green color aids in concealing it. When in the water, its cry 
is like that of B. lentiginosus americantts, but is more feeble, and very nasal" 
(E. D. Cope, Tex., 1893, p. 332). 

Breeding: Like B. debtlis, it doubtless breeds from mid-March to August. 
Ripe females have been secured to mid-July. 

Journal notes: Since the re-establishment of Bttfo insidior, we have had no 
opportunity to examine the collections of Bujo debilis and Bujo insidior to 
outline their contrasting characters or to determine their actual distinctness. 
The instant we saw Taylor's range of /?///<? insidior we thought of the time, 
July 10, 1925, near Fort Davis, Tex., when we heard narrow-mouthed toads 
and green toads; and also of the time in 1934 when we heard these same little 
toads at Alpine, Tex. On our 1942 trip we secured the toad near Balmorhca 
State Park. 

July 22, 1942, Toyahvale. Night trip west on highway. ... In this pond 
heard one lone Microhyla olivacea. Farther on heard plenty of Microhyla. 
Then I began to question whether they were all Microhyla. There was a call 
halfway between that of a Microhyla and a Bujo. A little toad under my feet 
put into the pond. I didn't see it well. Then I realized that it must be old Bujo 
debilis. Now that we were in the range of Taylor's Bujo insidior, might it be 



1 86 HANDBOOK OF FROGS AND TOADS 

that form? Caught three males (36, 58, and 43 mm.). As we came back heard 
many more, and where we heard it most, there on edge of road, saw a female 
just hit but yet alive. 

When we compared three Toyahvale specimens of B. insidior with three 
of our southern Texas B. debilis we seemingly had these differences: 

B. insidior B. debilis 

Snout 6.9-7.8 in L. 8.0-9.2 in L. 

Upper eyelid 9.5-10.0 in L. 11.5-12.5 in L. 

ist finger 7.6-8.6 in L. 8.8-11.5 in L. 

3rd finger 4-4-4-5 in L. 5.2-6.5 in L. 

4th finger 5.06-5.5 in L. 6.5-8.0 in L. 

Foot with tarsus 1.7-1.8 in L. 2.0-2.15 in L. 

Whether these measurements will obtain with a large series we dare not as- 
sert. They point to bigger snout, upper eyelid, first, second, and fourth fingers, 
and longer foot with tarsus in B. tnstdwr. 

Authorities' corner: "In September, 1886, coming down the valley of the 
Cimarron river from New Mexico, I first noticed the little Sonoran toad, Bufo 
debilis, Girard, near the Z H ranch in the Public Lands (now Beaver county, 
Oklahoma), at a point thirty-five or forty miles west of the southwest corner 
of Kansas. The species was observed a few days later in great abundance and 
activity (during rainy weather) in Morton county, Kansas, and in the south- 
ern part of Hamilton county. I have collected a single specimen in the western 
part of Barber county, Kansas, also" (F. W. Cragin, Kans., 1894, p. 39). 

"Two specimens (U.S.N.M. no. 2622) from Chihuahua, collected by Dr. 
Thomas H. Webb, are designated in the museum catalogue as the cotypes of 
Girard's Btifo insidior. The preservation of these specimens is fair, but both 
are very much bleached" (R. Kellogg, Gen., 1932, p. 52). 

When Dr. E. H. Taylor described a new species from a locality near Mazat- 
lan, he said that this small toad, Bufo folloggi (named in honor of Dr. Rem- 
ington Kellogg) "is most closely related to Bufo debilis and Bufo insidior 
Girard." "From Btifo insidior Girard (Kansas, Texas, New Mexico, Chihua- 
hua, Durango, and Zacatccas) it differs in the presence of the cranial crests 
(lacking or with only an occasional faint trace of crests in insidior), in having 
shorter hind legs, a shorter snout, larger eye, narrower head, and narrower 
interorbital width, a differently shaped parotoid and a totally different dorsal 
color pattern. It has very much larger and more numerous spiny tubercles on 
dorsal and lateral surfaces; the inner palmar tubercle is less developed, as are 
the metatarsal tubercles; the webbing between the toes is slightly more exten- 
sive. The color pattern is entirely different" (E. H. Taylor, Gen., i936a, 
p. 513). We have seen two B. f(elloggi and hold them distinctly different. 

"Bufo insidior Girard. The little green toad, although common in Texas 
and known from Kansas, has apparently not been recorded previously from 



TOADS: BUFONIDAE 187 

Oklahoma. Charles C. Smith and I found it to be fairly common in the 
southwestern part of Oklahoma where it was breeding with B. compactilis. 
Whereas B. compactilis was also found in great numbers on roadways, B. in- 
sidior was seen only in breeding pools. We collected specimens in Cotton, 
Jackson, southern Kiowa, and Tillman counties. In addition, Mr. Kuntz has 
furnished one specimen taken on a street during an evening shower in Law- 
ton, Comanche County; and the University of Oklahoma Museum of Zoology 
has a single specimen from extreme northern Garvin County" (A. N. Bragg, 
Okla., i94ia, pp. 51-52). 

"Under the name Bufo debihs, Campbell [Ariz., 1934, p. 3] recorded this 
species from the Huachuca Mountains in Arizona, and more recently Kauf- 
feld . . . has reported a specimen from the vicinity of Tombstone (30 miles 
south of Cochise). In view of the fact that Taylor . . . has reported speci- 
mens from Zacatecas, it is reasonable to suppose that the species occurs in 
northeastern Sonora. 

"The specimen collected by Kauffeld is now A.M.N.H. No. 50914 in the 
collection of the American Museum of Natural History. This specimen, an 
adult female with mature eggs in the oviducts, compares favorably in colora- 
tion and pattern with Zacatecan specimens depicted by Taylor . . . , particu- 
larly the individual depicted by him as figure 9 of plate 45. Taylor states that 
B. debilis, with which Kellogg (1932) synonymized insidior, inhabits Eastern 
Texas and Tamaulipas, whereas insidior occupies an extensive range from 
Kansas and New Mexico southward through Texas and the adjoining states 
in Mexico to Durango and Zacatecas. We have compared the Arizonan speci- 
men with individuals from Archer, Presidio, McLennan, and Bexar counties, 
Texas. Presumably some of these specimens would be referable to debilts, if 
Taylor's views are correct. Obviously there are differences in pattern and pos- 
tulation, but there is no conspicuous difference aside from these between the 
Arizonan and Texan specimens. The specimens from Archer County in 
northern Texas are certainly the same species as that from the other Texan 
counties, including Bexar, which lies immediately north of Atascosa County, 
where the toads depicted as debilis in figures 5 and 6 of Taylor's plate 45 were 
taken. If insidior can be removed from the synonymy of debilis, we strongly 
suspect that it represents a subspecies of the latter. Furthermore, if our speci- 
men from Bexar County is properly assigned to debtlis it is obvious that the 
form has a wider range than Taylor's statement implies, probably including 
Oklahoma' and Kansas, to judge only by a photograph of a Kansan specimen, 
and the proximity of Archer County to Oklahoma. San Diego County, Texas, 
where the toad in figure 4 of Taylor's plate was reputedly taken, seems to be 
non-existent; possibly the specimen came from San Diego, in Duval County. 

"Unless the type or other specimens from Chihuahua more closely resemble 
the Arizonan and Zacatecan individuals than Texan specimens, the applica- 
bility of the name insidior to the former is open to considerable question. 



1 88 HANDBOOK OF FROGS AND TOADS 

Taylor does not compare debilis with insidior, but compares each separately 
with kelloggi from Mazatlan. The latter evidently is a valid form inhabiting 
the coastal plain from Mazatlan at least as far south as Acaponeta in Nayarit 
where the specimens (A.M.N.H. Nos. 43877-43878) were secured by one of 
us in November, 1939" (C. M. Bogert and J. A. Oliver, Gen., 1945, pp. 409- 
410). See page 87 for Taylor and Smith's recent (1948) comparisons. 

Marine Toad, Giant Toad 

Bufo marinus (Linne). Plate XXXVIII; Map 10. 

Range: From southern Sonora to Tamaulipas in Mexico to Patagonia. In- 
troduced into Puerto Rico, Jamaica, and several smaller West Indian islands, 
some of which may possibly have had it originally native. Some distance north 
of William Lloyd's collection of it in Hidalgo, Tamaulipas, E. H. Taylor and 
J. S. Wright took it near Zapata, Tex. They wrote: "While collecting along 
the highway between Zapata and Arroyo Loma Blanca, in southern Texas, on 
the night of August 20, 1931, encountered a number of toads in a temporary 
pool, which on examination proved to be the widespread Bufo marinus 
(Linn.), the largest known species of Bufo." 

Habitat: In 1891 Ives found it in the courtyard of a house. In 1901 Waite in 
Bermuda reported them around houses, in gardens, and in water tanks. In 
1917 in Bermuda P. H. Pope stated that they were "abundant about roadsides, 
gardens and edge of mangrove swamps. A street light halfway between 'Gras- 
mere' and Hamilton was a favorite place for them, and two or three were 
usually seen under it, picking up the insects that were attracted by the light. 
... In the day-time they hide under stones or boards or burrow in the soft 
earth. I have sometimes seen them in little burrows in the side of a bank, 
where they had dug themselves in just enough to be out of the sun." In 1932, 
R. Kellogg declared that "these large toads are nocturnal in their habits and 
hide under fallen tree trunks, matted leaves, and stones, or burrow into loose 
soil." 

Size: One synonym, Bufo gigas Walbaum, suggests its size. It is the giant 
of toads. Gunther in 1858 (Gen.) remarked on the very large specimens and 
in 1885-1902 he noted that "Central- American specimens, at least those of the 
more temperate districts, do not appear to grow to the enormous size to which 
the species attains in Brazil and Guiana." His measured specimen is 166 mm. 
P. H. Pope (1917) in Bermuda wrote, "An average female measured 145 mm. 
from snout to vent and Waite describes one 155 mm. in length." "The male 
is smaller, usually about 13 mm. shorter, and more active than the female." 
Schmidt in 1928 had a i03-mm. female and a 95-mm. female from Puerto Rico. 
Kellogg (1932), among the 104 specimens he examined, mentioned individ- 
uals measuring no mm., 125 mm., and 167 mm. Taylor and Wright (1932) 
took several measuring from 66-168 mm., but the record seems to be that of 



TOADS: BUFONIDAE 189 

Miranda-Ribero, who reports a body length of 22 cm. (220 mm.) or 8.6 inches. 

General appearance: This is an enormous toad with an immense triangular 
pitted parotoid. The forward side of this triangular gland is just back of the 
tympanum; the base starts just back of the angle of the jaw and slants slightly 
upward to meet the very oblique or slanting upper side at a point well back on 
the side of the body. From this point a row or fold of large, blunt, rounded, 
brown-tipped warts extends % to % of the way to the groin. From Bitfo valli- 
ceps it may be distinguished by its very large parotoids; larger outer sole 
(metatarsal) tubercle; its coarser, less even, and less pointed tubercles; blunter 
and lower cranial crests; the usual absence or indistinctness of the parietal 
ridge; and less distinct color pattern. 

Color: We have seen none alive in recent years and consequently use Dr. 
Kellogg's (Gen., 1932) summary: "In general, the coloration of Bitfo marintts 
is quite variable, ranging through various shades of brown, including yellow- 
ish, reddish, and even blackish, and occasionally greenish olive; upper parts 
with or without large msuhform spots, which when present are usually edged 
with pale yellow; a light vertebral line occasionally visible; arms and legs of 
immature individuals usually banded with dark brown; underparts dingy 
white or yellow." 

Structure: Interorbital region smooth, decidedly concave, shallower in front 
than behind; crests pronounced the canthus rostrahs is a prominent crest 
beginning ahead of and above the nostrils and ending at the anterior corner 
of the eye, where it forks into two ridges, a broad preorbital and a well-defined 
supracihary crest which curves around the eye, sending off a prominent supra- 
tympanic ridge to the parotoid and a very short postorbital; parietal crests ab- 
sent, indistinct, or poorly developed; large broad head, box-snouted; parotoids 
huge, as long as head or bigger, % to i l /* times the head; parotoids widely di- 
vergent behind; toes M to % webbed; free inner metatarsal tubercle; outer 
metatarsal tubercle large and flat; a thin edged tarsal fold from inner meta- 
tarsal tubercle backward; prominent palmar pads at base of first and second 
fingers; eyelid finely tuberculate; two rows of large fleshy warts, one down 
each side of middorsum to vent, more prominent in male. Back of angle of 
mouth the skin is divided into two or more vertical folds so that at times it 
might be thought a postrictal gland. However, it is not like the white wart of 
Bufo alvarius. 

Voice: Taylor and Wright heard these toads at Zapata, Tex., but did not 
describe their call. Pope in Bermuda has stated that they have a more resonant 
and louder call than Bufo amertcanus "a deep booming trill." 

Breeding: In Bermuda February to July is the breeding season, April being 
the optimum month. In Trinidad they breed from August to October; in 
Demerara, from mid-April to September, possibly also from November to 
January. The eggs are in strings. Dr. E. L. Mark had some hatch in 68 hours. 
The black tadpole transforms after 45 days or less. The tops of the first two 



190 HANDBOOK OF FROGS AND TOADS 

fingers of the males have excrescences, and there are excrescences on the inner 
side of third finger and the inner palmar tubercle. Some males have brown 
and spiny-tipped warts and tubercles. 

Clark has a note that this species lays twice a year, but Pope is sure that in 
Bermuda an individual breeds but once a year. In Bermuda in 1903 Mark 
made the following observations : 

"In the spring of 1903 ... we visited 'Spanish Rock' near midday, April 
22. Before reaching the Rock we found, on the slope facing away from the 
ocean and toward Spittal Pond, a small pool of rainwater (there had been a 
heavy shower the night before), and in this pool there were large numbers 
of the huge toads Bit jo aqua in pairs. The females were engaged in spawn- 
ing, and the numerous strings of spawn were stretched across the pool in 
almost every conceivable direction. The pool, some fifteen or twenty feet in 
diameter and only a few inches deep, was of so temporary a character that its 
bottom consisted throughout of turf, not unlike the land immediately sur- 
rounding it, which was not submerged." 

Ruthven (1916) "recorded that on the Demerara River, about thirty-five 
miles south of Georgetown m 1914, tadpoles were abundant in July and Au- 
gust. . . ." Pope (1917) thought that development might be more rapid than 
Ruthven's 45 days from egg to adult. Mark (1903) observed this species from 
eggs April 22 to free-swimming larvae April 27. In describing some tadpoles 
collected by L. J. Cole in Bermuda Pope wrote: "The tadpoles are black in 
color and resemble those of B. americantts. They measure from 8.5 to 10.9 mm. 
in length. They have the typical early tadpole form. The buds of the hind 
limbs are present, but hardly visible to the naked eye." 

Transformation must be over as extended a period as is the egg laying. 
Ruthven records it on August 16 about 45 or 46 days after egg laying. No 
other notes apparently exist on transformation. 

Journal notes: We present none. Our experiences consist in seeing a few 
captive specimens in aquaria or zoos. 

Authorities 9 corner: Taylor and Wright (Tex., 1952, pp. 247-249) found 
their specimens had been feeding on "tenebriomd and carabid beetles, the 
bulk of the food being of the genera Eleodes and Pasimachus." Pope (1917) 
says that when the residents of Bermuda found that these toads ate cock- 
roaches, they no longer called them after their introducer "Captain Vesey's 
nuisances." In 1918 Noble found that the Bufo marinus he had from Nica- 
ragua fed principally on cockroaches. Our friends, Drs. M. L. Leonard and 
Stuart Danforth, years ago (1930 or earlier) told us how some of their co- 
laborers in Puerto Rico had successfully shipped them to Hawaii to control 
insects, and today plenty of reports appear on their role in Hawaii. 

Regarding the poison Vernll says: "This toad is believed ... to have a 
very poisonous secretion from its parotoid and dorsal glands. It is said that 



TOADS: BUFONIDAE 191 

dogs that mouth them invariably die within a few hours. The secretion of the 
glands, when injected into the circulation of dogs, birds, and other animals, 
causes convulsions and death, even when in small doses. Mr. A. H. Verrill 
... on one occasion saw the venom ejected as a fine spray, from the parotoid 
glands of a large toad, when it was much irritated." 

On this topic the observations of C. T. Dodds seem pertinent: "During 
the summer of 1922 I spent some time at Los Mochis, in Northern Sinaloa, 
Mexico. It was very interesting each evening to note the appearance of 
large toads under the electric lights. One evening, about the end of July 
while one of these toads was occupied in catching insects, a small terrier dog 
started to tease the amphibian. At first the toad only hopped in an effort to 
escape the barking dog, which did not attempt to bite. Soon the toad became 
tired and its hops were less frequent and not so long. The dog was urged on 
by the bystanders and finally snapped the toad while the latter was in mid- 
air, catching it just back of the foreleg. Although the dog's mouth was not in 
contact with the toad for more than an instant, he immediately lost all inter- 
est in the animal. Spitting and shaking his head, he gave all indications of 
having received something very distasteful m his mouth. He was offered 
some water but refused to drink and in about a minute's time showed signs 
of weakening and general paralysis. He sank to the ground with his legs 
spread out, writhing and whining with pain, and unable to recognize his 
master. During this time he was able to push himself along the ground, grad- 
ually becoming weaker and very rigid, with eyes greatly protruding and 
respiration and heart action exceedingly rapid. After twenty minutes he was 
somewhat quieter as if he was going to die. It was suggested that castor oil 
be given him in the hope that it might save his life. Accordingly his mouth 
was pried open, for every muscle was rigid, and about 50 cc. of oil was poured 
down his throat. Ten minutes later he showed signs of improvement, al- 
though I do not know that the oil had anything to do with the change. Within 
another 15 minutes he was able to get upon his feet but was very feeble and 
his hind quarters were still somewhat paralyzed. Presently he recognized his 
master and within an hour from the time he took his distasteful nip he was 
apparently quite normal again." 

E. H. Taylor and H. M. Smith recently expressed the following opinion: 
"We are convinced that Bitfo marinus Linnaeus, as generally accepted, 
comprises species or/and subspecies and is in fact of almost generic signifi- 
cance. However, certain difficulties are involved in properly delineating and 
naming these forms. In the first place the type localities of Bufo marines and 
B. aqua are unknown except that they are from the Western Hemisphere. 
Bufo maculiventris and B. lazarus of Spix are Brazilian, but lack exact locali- 
ties; B. ictericus Spix, however, is cited with Rio de Janeiro as type locality. 
B. htimeralis Daudin 'existe dans diverses countries meridional du nouveau 



192 HANDBOOK OF FROGS AND TOADS 

continent.' He mentions one in the Musee d'Histoire naturelle from Cayenne 
(French Guiana). This may be presumed to be the type locality for this 
species. 

"A second difficulty is that seldom are good series of these great toads col- 
lected; and the age, sex, and environmental variations are known for only a 
few localities. 

"Wiegmann described Bufo horribilis from a series of cotypes from the 
State of Vera Cruz, and we are reviving this designation for most of the toads 
of this group in Mexico, aware that there are probably variant populations 
even here, that may warrant subspecific designations' 1 (Gen., 1945, pp. 551- 
552). 

"The third series of eggs was at the yolk plug stage when placed in 15 and 
10% sea water and in tap water. Mortality appeared to be slightly less in the 
dilute sea water, and development in the early stages, at least, somewhat ac- 
celerated. 

"Twenty-seven days after laying, the first completely metamorphosed ani- 
mal was taken from the 15% sea water and 2 days later metamorphosis was 
occurring in the other solutions. Thereafter, over a period of 9 days, meta- 
morphosed animals were taken from all solutions. 

"In general, these preliminary experiments seem to show that low concen- 
trations of sea water constitute a favorable environment for the development 
of Bufo marintis larvae" (C. A. Ely, Gen., 1945, p. 256). 

"In the latter part of November, 1939, the senior author secured several 
specimens of Bufo marinus under boulders along the river at Culiacan in 
Sinaloa. This was during the dry season and apparently the toads were hiber- 
nating. While these specimens were being collected a native boy watched with 
interest as the toads were placed in the sack and finally commented, 'Echnn 
leche' (they throw milk), in reference to the whitish venom secreted by the 
parotoid glands. This secretion is readily ejected a distance of a foot or so if the 
glands are squeezed. However, the toads have not been observed to expel the 
poison such a distance voluntarily. 

"It has been noted on several occasions that B. marmits, when killed by 
being placed in formalin or alcohol, commonly secretes quantities of viscous^, 
whitish liquid from the parotoid glands" (C. M. Bogert and J. A. Oliver, 
Gen., 1945, pp. 340-^41). 

Spotted Toad, Belding's Toad, Canyon Toad, Red-spotted Toad 

Bufo punctattis Baircl and Girard. Plate XXXIX; Map 13. 

Range: South-central Texas west to Nevada, southern California, Sonora, 
and Lower California. 

Habitat: Desert canyons, breeding in rock-bottomed pools of intermittent 
streams. 



TOADS: BUFONIDAE 



193 




Plate XXXVIll. Bufo marinus. 1,2. Male Plate XXXIX. Bufo functatus. i. Male 

(X%). 3,4- Male (X%). croaking (X%)- 2,3,5- Males (X%). 4- 

Female (X%). 



I 9 4 HANDBOOK OF FROGS AND TOADS 

"Lower California, Aguaito Springs, 15 miles E. Rosario, 1300 ft. lat. 304'; 
37244; June 9. 2 miles NNW Catavina, 1950 feet, lat. 2947 / ; 37245-37248; 
June ii. 

"The red-spotted toad from Aguaito Springs was the only individual of 
this species seen at that locality a series of spring-fed pools in an arid terrain. 
Of the four from near Catavina, two were hopping at night on the dry, sandy 
floor of an arroyo at least 100 feet from water, and two were squatting in a 
pool" (L. Tevis, Jr., L.C., 1944, P- 6). 

Size: Adults, i %-$ inches. Males, 40-68 mm. Females, 42-64 mm., or even 
74 mm. in Lower California. 

General appearance: Tins is a small, delicately formed, alert, attractive toad 
of grayish, greenish tan, ttiupe, drab, or even red color, with a flattened body 
and a broad flat back evenly covered with scattered tubercles of small size. 
The tubercles on the back, sides, and legs may be reddish, orange-cinnamon, 
or light vinaceous-cmnamon. There may be black rings or partial rings at 
the bases of the tubercles. The under parts are buff or white and may be 
spotted with black in the smaller toads. The legs may be barred or spotted 
with black. The conspicuous marks of this toad are the small, round paro- 
toids, the broad interorbital area, and the sharp-edged, often pebbly, canthus 
rostralis that gives the nostril a "boxed" appearance. The eyelids also are so tu- 
bercular as to appear pebbly. It is a fine little toad of very neat, compact ap- 
pearance. It frequently gives a pleasant birdhke chirp in captivity. We picked 
up one and turned it over. Ir lay m the shallow water with legs drawn up as 
if to "possum"; we never saw any Bnfo feign this lifeless attitude more than 
this individual. 

Color: Female. Hclotes, Tex., March 1 3, 1925. Upper parts light brownish 
olive,' bufTy olive, or buffy brown on parotoids and mterparotoid area. (The 
next 'day the upper parts in general were drab.) Upper part of hind legs dull 
citrine sharply marked from light under parts by wax yellow or primulme 
yellow. This color or cream color on underside of thighs. Tubercles on top, 
lateral, and ventral surfaces of thighs and on underside of fore and hind feet 
are orange-cinnamon or vinaccous-cinnamon to light vmaceous-cinnamon. 
Under parts cartridge buff, the area ahead of hind legs olive-gray. The ins has 
cream or pale chalcedony yellow rim around the pupil. It is black heavily" 
spotted with tilleul buff, vinaceous-buff, and light vinaceous-fawn. 

Male. Grand Canyon, Oct. 12, 1929. Back buffy citrine or olive-lake, buffy 
olive, citrine-drab, or light grayish olive. Legs fore and hind olive-buff. Over 
back are scattered little spots of flame scarlet, rufous, or apricot orange. On 
sides are several spots of black, some without and some with rufous or apri- 
cot orange centers. On back each spot has such a center. Black or grayish 
around snout to eye, more or less around tympanum, slightly on upper eye- 
lid, and somewhat on parotoid. Forelegs spotted with black but these spots 



TOADS: BUFONIDAE 



'95 



with centers of ground color. Femur, tibia, tarsus, and outer toe with bars of 
black. Upper labial margin olive-buff. Undersides of hands and feet and more 
or less of fore limb and hind limb cinnamon or cinnamon-buff. Throat 
honey yellow, old gold, or tawny-olive. Lower throat with gnaphalium green, 
tea green, dark bluish glaucous, or greenish glaucous-blue. Lower belly and 
most of buttocks light brownish drab, army brown, or deep brownish vina- 
ceous. Eyelids above and canthus heavily tubercled with rufous or apricot 
orange. Iris rim broken above and below, behind and in front viridine green; 
iris black, heavily dotted and splashed with ochraceous-salmon, ochraceous- 
buff, or antimony yellow. Under parts white dotted with small black dots. 




Map 13 

Another male has no spots underneath. A third has them on forward chin 
and rear pectoral region, those on chin white-centered; a fourth has them 
faint on pectoral region. One is light drab, another cinnamon; the drab one is 
avellaneous or wood brown, uniform on back except for reddish tubercles. 

Specimen collected by V. Bailey, at the bottom of the Grand Canyon, Sept. 
13, 1889. "I took the following color description from one of the fresh speci- 
mens (No. 16185); Above 'malachite-green' densely speckled with small dots 
of bright vermilion; limbs paler, dotted with vermilion and also with minute 
black specks which likewise occur on the flanks; region surrounding nostrils 
black; upper lips and whole under surface bluish white, irregularly speckled 
with black; posterior part of belly and underside of thighs dark brownish flesh 
color; soles, dull orange" (L. Stejneger, Ariz., 1890, p. 117). 

Structure: No cranial crests, or crests obscure; parotoids small and round; 



196 HANDBOOK OF FROGS AND TOADS 

interorbital area broad; ridge from nostril to eye sharp and often pebbly, giv- 
ing the nostril a "boxed" appearance; finger excrescences of males not promi- 
nent; throat of male somewhat discolored. 

Voice: The vocal sac of the male is a round subgular pouch. The call is 
birdlike. It is a high-pitched, yet very pleasing, trill, lasting while one counts 
to 15 or 25. 

"Cape Region of Lower California. While collecting with a light early in 
the evening forty-six specimens were taken around the public square in the 
little village of San Antonio. They were heard calling late in the evening. 
A specimen captured was observed to make a shrill whistling noise of four or 
five seconds duration, at intervals of about the same length" (J. R. Slevin, 
B.C., 1928, pp. 101-102). 

"A long continued clear trill, resembling that of a hearth cricket but with 
more volume" (J. Grinnell, MS; and T. I. Storer, Calif., 1925, p. 197). 

Breeding: They breed from April to September, May being the most com- 
mon month. The eggs arc single, with very sticky jelly, and sometimes the 
eggs are stuck together loosely as a small film on the bottom. The envelope is 
single % inch (3.2-3.6 mm.), the black and white vitellus M>rr-%o inch (i.o- 
1.3 mm.). The eggs hatch in 3 days or less. The small tadpole, i inch (24-25 
mm.), has a very black body and a whitish tail with many evenly spaced black 
dots. The tooth ridges are %. After 40 to 60 days, the tadpoles transform from 
June to August, at % inch (9-11 mm.). 

Journal notes: July 19, 1917, from 5-7 P.M. at Texas Pass, Ariz. We heard 
plenty of toads but could not find them. At last P. A. Munz and I dug a 
croaking male from beneath a rock. They croak in the crevices and under- 
neath rocks. In the pools are plenty of tadpoles large and small. In the evening 
we picked up a female B. punctatus. In the canyon were no end of males in 
the water and along the banks, males seizing each other. As yet there are few 
females. 

Aug. 2. We started down Bright Angel Trail at Grand Canyon. At Indian 
Gardens, in a flat shallow area, water 6 inches deep, we found two egg com- 
plements of B. pnnctatus on the muddy bottom. They were near the west 
edge in shallow water and not under overhanging willows nor in the water 
cress. The eggs were more or less agglutinated on the bottom about one egg* 
deep, in a few places more. Later, we found many complements in all stages 
and took three or four transformed stages of Bufo punctatits here. 

May 6, 1925, Helotes, Tex. Where I found the female B. punctattis and two 
males, there are strewn over the bottom eggs, black and white. Some are 
single; some touch in masses but not strings or masses which would stay as 
masses if above the bottom. These occupy a foot square or i by i^ feet. These 
are nearly hatched. On the rocky bottom they look like the black-fly masses 
we get in swift water in the Northeast. Water here is i inch deep. Nothing but 
B. pttnctatus was calling here last night or the night before. We discovered 



TOADS: BUFONIDAE 197 

these at 11:30 A.M. We photographed them at i P.M. Just now under some 
near-by stone a B. pttnctattts male can be heard. 

June 3, 1942, St. George, Utah. Went up on Red Hill near Dixie sign. In a 
ditch Anna saw tads and some eggs. She found a dead B. functatus in ditch. 
Under a stone I took a Bufo punctattts. 

Authorities' corner: 
G. P. Englehardt, Ariz., 1917, p. 6. 
N. N. Dodge, Ariz., 1938, p. 12. 
R. R. Miller, Calif., 1944, p. 123. 

Oak Toad, Oak Frog, Dwarf Toad 

Bufo quercicus Holbrook. Plate XL; Map 14. 

Range: North Carolina to Florida west to Louisiana. 

Habitat: Abundant in the pine barrens, seeking shelter in little burrows 
shielded by vegetation or under boards or logs. Many breed in shallow cypress 
ponds or in temporary surface rain pools or ditches. 

Size: Adults, : J1-iJ4 inches. Males, 19-30 mm. Females, 20.5-32 mm. 

General appearance: This pigmy toad has a light stripe down the back and 
four or five pairs of unconnected spots along the middle of back, from the 
first pair between the eyes to the last, which are merely two pinpoints just 
ahead of vent. They may be light brown or almost black so that the spots 
barely show. The skin is finely roughened with tubercles, many of which are 
red. This brightly colored little fellow looks like a bit of velvet or tapestry. 
The arms and legs arc barred with black. The vocal sac of the male is con- 
spicuous when deflated, and is a triangular apron with the base on the gular 
line and the point extending backward over the pectoral region. Under parts 
grayish or buffy. 

Color: June 8, 1921. Adult. Dorsal stripe white, pale orange-yellow, maize 
yellow, sulphur yellow, or cream-buff. The four or five pairs of dorsal spots are 
black. Upper parts with some gull gray, pearl, or pale olive-gray. Stripe from 
lower part of tympanum almost to groin of the above grays; also patch of the 
same back of angle of mouth, below tympanum, above arm insertion, in front 
of femur, and on back part of upper eyelid. All these lighter dorsal portions 
are with burnt sienna tubercles, which are especially prominent along either 
side of dorsal stripe from hump backward, on oblique lateral stripe, and on 
posterior part of eyelid. The tubercles on black areas look black but many are 
really burnt sienna. Parotoid with fine and thickly studded burnt sienna tu- 
bercles. Tubercles on palmar and solar surfaces, posterior surface of thighs 
(partially), groin (a little), pectoral region (a few tubercles) vinaceous- 
rufous, Hay's russet, or mars orange. Ventral parts smoke gray, grayish white, 
pale olive-buff, or cream color. Each tubercle stands out: on venter, close to- 
gether and black between; on sides and underside of limbs, wider apart and 



198 HANDBOOK OF FROGS AND TOADS 

intervening black more apparent. Iris cream color; rim around pupil and eye 
naphthalene yellow; rest largely black with cartridge buff or ivory yellow. 

Structure: Head to angle of mouth short; snout pointed; body short; flat; 
hind limbs shorter than body length; first finger less than or equal to second; 
cranial crests divergent, ends connected by transverse series of raised warts, 
giving the cranial hollow a parapet behind; parotoids finely spmose; excres- 
cences on fingers of male not prominent; interorbital region broad. 

Voice: The vocal sac of the male is an oblong "sausage." When deflated it 
is made up of folds ot skin on the lower throat covered by a conspicuous apron 
or lappet. The call is very birdhke, not f roghke. It is a very high-pitched whis- 
tle. The chorus is deafening and can be heard % mile or more. 

May 16, 1921. The oak toad male was calling before we approached. He 
piped only low. After we had worked him around for a photo, he suddenly, 
to our surprise, backed into a hole at the base of a saw palmetto. The hole was 
% of an inch in diameter and not deep. The note of this toad is birdhke. One 
will hear three or four calls sounding like those of piping chickens. Sometimes 
the note is repeated three or four times. Then this process is repeated after a 
short interval. Once us note was likened to that of a swallow-tailed kite. Truly 
the most unfroghke note 1 ever heard. It sometimes sounds like some animal 
in distress. There are several calling in the pmey woods. One calling from 
a tangle of chokebernes (Aronta), Qsmttnda cinnamotnea, Bamboo brier 
(Smilax), and sweet bays. Couldn't find it. Are the toads moving pondward? 
Later in the evening we heard none. 

1922. A combined chorus of Hyla gratiosa, Hyla femoralts, Hyla sqtttrclla, 
Chorophiltts, Acns, and Bitfo qucrcicits left our ears ringing for a long time 
after we left the pond. By half closing our ears, we shut out Hyla jemoralts 
and heard the others more distinctly. The whole was a terrible din. 

In 1912 and 1921 some of the residents who were almost invariably accurate 
assured me that the black snake had a whistle and that this note of the oak 
toad was the call in question. The timed calls ranged from $2 calls in 16 seconds 
to 10 calls in 26 seconds. 

In inflation of the throat these toads are like Kt/fo comfactilu and Bujo cog- 
natus. The lower throat is the principal part involved in the process; it is 
thrown out into an elliptical bag or sausagelike balloon. One can tell when 
a toad is going to trill after a rest because the body will inflate to a large size 
and then the ludicrous sac projects in the throat region. The tip of the sac 
when not really inflated comes close to the tip of the chin. Otherwise it ap- 
pears as a little, loose, vibrating sac % cm. out from the lower part of the 
throat. When the sac is deflated, the body inflates. When the body is com- 
pressed or deflated, the sac inflates. 

Breeding: They breed from April i to September 5. It takes a heavy warm 
rain to start these little toads calling vigorously. The eggs are in bars of 2, 3, 4, 
5, or 6, and the bars are from Vrz-V* inch (2-7 mm.) long and %o inch (1.3 



TOADS: BUFONIDAE 199 

mm.) wide. The eggs, black and white, },- inch (i mm.) in diameter, are laid 
on the bottom of shallow pools. The small tadpole is grayish with six or seven 
black saddles on the musculature, and with heavily marked upper tail crest, 
and the venter is one mass of pale purplish vinaceous. They transform July 13 
to August 1 6, at ^d-'Ke inch (7-8 mm.). 

Journal notes: May 26, 1921, Billy's Island, Okefinokee Swamp. In pipe- 
wort, sedge, and grassy places at 10 A.M. found a female Eu]o qtiercicus. We 
hear males in the woods. The calls are more lively and insistent. Is B. querci- 
ciis going to the ponds soon? We have taken four or five toads this morning. 
Females are about more since last night's thunderstorm. In a burnt-over area 
it seemed as if more were present. Possibly they are easier to find in this area. 
We found three males and three females. The males arc not in holes. 

June 4, Billy's Island, Okefinokee Swamp. About two inches of rain 
dropped, and the island seemed teeming with oak toads. They bred almost 
everywhere. All about the cleared fields, in piney woods, in hammocks, and 
in numerous other places, we found oak toads that day. On July 3 the species 
was abroad in great numbers. Every transient shallow pool filled by the rain 
had them calling. We took three or four pairs and 30 to 40 males in short 
order. 

On July 27, 1922, in a shallow pond, we heard so many oak toads we looked 
for eggs. We found single bars of two to six or eight eggs rarely attached to 
sticks at the surface, usually attached to grass blades 0.5 to i or 2 inches below 
the surface of the water, the water i to 3 inches deep. Other bars were attached 
to pine needles. Once in a while two bars extended out from a common focus. 
Normally they were close together. 

Authorities' corner: 

J. E. Holbrook, Gen., 1842, p. 14. P. Viosca, Jr., La., 19233, p. 37. 

E. Loennbcrg, Fla., 1895, p. 338. A. F. Carr, Fla., i94ob, p. 54. 

H. W. Fowler, Fla., 1906, p. 109. 

Southern Toad, Carolina Toad, Gray Toad, Land-Frog (Bartram), Land- 
Toad (Catesby), Latreille's Toad, Charming Toad, Hop Toad 

Bujo terrestns (Bonnaterre). Plate XLI; Map 9. 

Range: North Carolina to Florida, west to the Mississippi River. We rather 
incline to the limitation of this form to the coast below the fall line. Are the 
isolated records (one, western South Carolina; one, northern Georgia; one, 
northern Mississippi; one, northern Louisiana; several, extreme western 
Louisiana) misidentifications or intermediates? S. N. Rhoads in southeastern 
and middle-central Tennessee calls his specimens intermediate. 

Habitat: Abundant throughout its range, particularly common in culti- 
vated fields. It occurs throughout the pine barrens and hammocks, in fact in 
any land habitat. When breeding, it is usually in shallow water from the tiny 



166 



HANDBOOK OF FROGS AND TOADS 








Plate XL. Bujo quercicus. 1,2. Males 
croaking (X%)- 3.8- Males (XO- 4,6.7.9- 
Females (XO- 5- Egg bars 



Plate XU. Bufo terrestrit. i. Mai 
(X%)- 2. Male (X%)- M- M al croak 
ing (X%). 5- Female (X%). 



TOADS: BUFONIDAE 201 

pool to the edges of lakes. Frequently these toads are in pools so transient that 
they can last but a few hours, as in the furrows in fields or in temporarily over- 
flowed grassy areas. On one occasion their eggs were so plentiful in imperma- 
nent pools that we wrote (June 4) : What a frightful waste of frog life in tran- 
sient pools! 

Size: Adults, 1^-3% inches. Males, 42-82 mm. Females, 44-92 mm. 

General appearance: These toads vary in color from red or gray to black. 
The crests on the head are prominently raised at the rear into clublike promi- 
nences or knobs. The skin between the larger warts is finely and uniformly 
roughened with tubercles all over, including the eyelids and parotoids. 

Color: June 9, 1921, Okefinokee Swamp, Ga. Male. Line from front line of 
eyes almost to vent pale gull gray or mineral gray. Black spots along dorsum: 
one pair near cephalic edge of eye; one pair from upper eyelid connected 
across the meson; one in the middorsum between tympana; a pair either side 
of middle between the rear ends of tympana; a pair of small spots; a pair of 
large spots where hump comes; a pair of small spots; a pair just ahead of vent 
all black spots of dorsum thinly encircled with chalcedony yellow. More 
or less broken pale gull gray or mineral gray line from tympanum to groin; 
below this a prominent black area. Tubercles of back black or deep brown 
tipped. Lighter areas on rear of hind legs sulphur yellow. Either side of vent 
a few orange tubercles. Under parts pale smoke gray. Pectoral region, under 
parts of hind legs, and sides with black spots. Lower jaw rim like belly color. 
Throat deep mouse gray or dark mouse gray with widely spaced white dots, 
giving throat discolored appearance. Light area of dorsum of hind foot same 
color as the rear of thighs. Very little rusty on front of thighs and groin. Top 
of first two fingers with excrescences and slight line of such on edge of third 
finger. Color of excrescences chocolate, or better, Hay's maroon or maroon. 

Female, Lighter, larger. Practically no spots on pectoral region. None on 
throat. Throat same color as belly. Practically no rusty spots. Ground color of 
the dorsum more greenish olive. Sometimes at breeding season males and 
females may be alike in color, e.g., on April 24, 1921, several were thus, several 
pairs reddish, one pair gray. Most of the pairs, however, were diverse. 

Structure: Prominent knobhkc crests; backs of thumb and second finger 
of male with excrescences, which are also on inner edge of the third finger; 
female throat usually light; male throat usually dark; pectoral region may be 
heavily spotted, sometimes only a median spot. 

Voice: The note sounds much like our droning Bufo americantts of the 
North, not like the scream of Bufo w. jowleri. The trill is perhaps shorter than 
that of B. amcricanus. The trill lasts 7-9 seconds, with intervals of 4-60 sec- 
onds. It is musical in character. It has been described as a high trill, a drone, 
or even a bass roar, for when many are calling close to the observer, the sound 
is deafening. The choruses can be heard some distance away. Like other 
species, they may give weak notes; individuals may be freakish, hesitant, 



202 HANDBOOK OF FROGS AND TOADS 

shrill, or even, rarely, open the mouth to scream, or with half-inflated throat 
give puzzling notes. Usually when croaking, the throat is distended to its full 
capacity with the body compressed. Then the body is distended and the throat 
collapsed. 

The calling toads in cypress ponds and bays may be perched on a log, on a 
cypress knee, stub, or stump, at the base of a cypress or gum tree, on the moss, 
or resting on aquatic plant stems, leaves, or dead twigs, usually in shallow 
water or at the edge of a pond. In overflow pools they may be anywhere in 
shallow places. Rarely if ever do they float when croaking. This toad is truly 
an alert, pert animal. 

Breeding: These toads breed from March i, or earlier, to September. The 
eggs are in long coils of jelly, the egg !4v-Vi<i inch (1-1.4 mm.), the outer 
tube Mo-%6 inch (2.6-4.6 mm.), the inner tube Mu-% inch (2.2-3.4 mm -)- 
The eggs, separated in the tube and with no partition apparent between 
them, number 2500-3000, and hatch in 2-4 days. In the small tadpole, i inch 
(26 mm.), the body is broader toward the rear, the tail crests are narrow, the 
tail is short and rounded, and the eyes are dorsal, close together; it is black in 
color. The tooth ridges are %. After 30-55 days, the tadpoles transform from 
April to October, at /4-% inch (6.5-1 1 mm.). 

Journal notes: April 24, 1921, Okefinokee Swamp, Ga. At 9:15 A.M. Bitfo 
terrestris were calling. Water 66. Sun very bright. There were 15 pairs in an 
area 6 feet square. One pair, both male and female reddish. Male about % 
size of female. Male embraces female in axillary fashion, the last two fingers 
not dug into axil but resting on arm insertion of the female. When the pair are 
not laying, the male has the hind legs free and floating, but when the female 
ovulates several inches of egg string, the male brings his knees into the groin 
of the female and heels almost touch, the upper surface of hind feet against 
the underside of femur and near cloacal opening. Female with hind legs 
stretched back sometimes heels touching, sometimes not. The eggs rest in the 
cup made by heels and feet of male. The pair may remain in emission atti- 
tude 4 or 5 minutes or less. Then the female crawls i foot or more. A minute 
may elapse before another emission. By 11:30 A.M. surface of water was 81. 

May 2. One of the boys called this toad "charming toad," because "it charms 
you, turns your eyes right green." 

June 8, 1930, 2 miles north of Mandeville, La. In the pine-barren parishes of 
eastern Louisiana, Viosca finds Bttjo terrestns; the first one I espied in a mixed 
magnolia thicket was as intense a reddish brown as I have ever seen in a toad. 

Authorities 1 corner: 
J. E. Holbrook, Gen., 1842, 5, 9. 
R. F. Deckert, Fla., 1914, p. 2. 



TOADS: BUFONIDAE 203 

Nebulous Toad, Wiegmann's Toad, Mexican Toad 

Bufo valhceps Wiegmann. Plate XLII; Map 13. 

Range: Louisiana, eastern and south-central Texas to Mexico and Costa 
Rica. 

Habitat: Lowlands in the West Gulf Coastal Plain of Louisiana, in the hills 
in northwestern Louisiana, in the pine lands of eastern Texas, and in open 
stretches of streams in central and southern Texas. Frequently found in rail- 
road ditches or roadside pools. 

Size: Adults, 2^-5 inches. Males, 53-98 mm. Females, 54-125 mm. 

General appearance: This is a large brown toad, with a light streak down 
the middle of back from snout to vent, a light area over each parotoid extend- 
ing diagonally backward to the groin. This light lateral area is bordered below 
with a fringe of white conical tubercles. The skin of back and venter is closely 
set with tubercles. There is a light line on the upper lip, continued beyond the 
rear of the angle of mouth by another row of light tubercles. The throat of 
the male is discolored a citrine drab or water green. 

Color: Male. Beeville, Tex., March 12, 1925. Parotoid and interparotoid area 
wood brown, fawn color, or cinnamon, changing to drab or buffy olive on m- 
terorbital and upper eyelid, face, and nasal area. Stripe down middle of back 
deep olive-buff, vmaceous-buff, or pale pinkish cinnamon. Stripes on either 
side the same color. Stripe on upper jaw tilleul buff or cartridge buff. Light 
spots on lower side the same color. Belly cartridge buff becoming from pec- 
toral region rearward cream-buff, spotted with black. Dark bands on side and 
back brownish olive. The dark lateral band may be clove brown on side and 
olive-brown above. Two pairs of black spots on each side of dorsal stripe. 
Hind legs and forearms banded with deep olive, the interspaces wood brown 
or avellaneous. Front of chin cream-buff. Pectoral region pale vinaceous-fawn, 
rear chin discolored citrine-drab or water green. Iris pupil rim ivory yellow or 
pale green-yellow with a warm sepia line above it; iris largely tawny, ochra- 
ceous-orange, or lemon chrome; lower eyelid transparent with spinach or calla 
green rim; rest of lower eyelid with scattered spots of this green. 

Female. The female has dorsal stripe and light areas pale olive-buff, while 
two males have them dark olive-buff. Top of head and parotoids grayish olive 
or light grayish olive. Upper parts olive or deep olive with little of brownish 
shades as in males. Under parts including throat deep olive-buff, cleaner on 
throat than on abdomen, where it is rather dirty in appearance. 

April 2, 1925, Nueces River, Tex. Found a young B. valhceps under a log. 
It has about a dozen scattered, prominent, buff, yellow, or light cadmium spots 
on the back. Very spotted under parts and particularly in midpectoral region. 

Structure: High projecting crests on crown of broad head; these are canthal, 
preorbital, supraorbital, postorbital, parietal, and orbitotympanic ridges; paro- 



204 HANDBOOK OF FROGS AND TOADS 

toids rather small, round, or triangular; snout obtuse; toes % webbed; inter- 
orbital space wide; internasal space narrow; upper eyelid much less than 
interorbital space; male with a subgular vocal sac not revealed by wrinkles on 
the throat; body flat; mouth large; excrescences on two fingers of male promi- 
nent. 

Voice: The vocal sac is a large, round, subgular pouch. The call is louder, 
harsher, and lower in pitch than that of Bttfo americanus. The croak lasts 3-4 
seconds. Often the males take stands several feet up from the pond's edge. 

March 24-26, Beeville. Bufo valliceps calling vigorously along roadside 
ditches, in streams and around tanks near windmills. Found no females or 
mated pairs. Night of March 25. Bujo valliceps males are hard to photograph 
at night. After all is set up, they usually move when the flashlight is taken off 
preparatory to the gun flash. What a time between the moving toads and the 
frequent cars! This was a roadside pool. 

March 28, San Diego-Alice, Tex. By railroad and creek bridge heard Bujo 
valliceps. At first thought it was a toad much like our B. americanus. It 
sounded something like it at a distance, and not harsh as it is when near by. 

Breeding: This species breeds from March to August. The eggs are often 
in double rows in long strings of jelly with the wall of the inner tube close to 
the outer. This jelly grows looser with age, so that there may be a double row 
of 25 to 27 eggs in j% inches (3 cm.) or a single row of 7 to 10 eggs in i% 
inches (3 cm.). The outer tube is Y& inch (3 mm.), the inner tube M.O inch 
(2.6 mm.), and the vitellus %o inch (1.2 mm.). The eggs are purplish black 
and hatch in iVz-2 days. The small blackish tadpole has 8-10 black bars with 
intervening pale buff areas on the dorsum of the tail musculature. The tooth 
ridges are %. After 20-30 days, the tadpoles transform, April to September, 
at Ar-^A inch (7.5-12 mm.). The season varies with the periods of heavy local 
rainfall from March-August. Pope found a female ready to spawn as late as 
August 25 in Houston, Tex. 

May 10, 1925, Helotes, Tex. Captive pair laid eggs. Egg strings. Some of the 
eggs seem to be in double rows in string; other parts of same string may be 
with single rows. There are many single eggs or eggs without string effect. 

May 13, Helotes. Went to Marnock's Second Crossing. Found B. valliceps 
egg strings widely spread out. These were long strings in midwater and buoy- 
ant in midplane. They were more above the bottom than eggs of any toad I 
have seen. One bow of two strings, 3 feet long, was attached only at ends. 
These eggs covered an area of 6 or 4% feet square. They seem to be the com- 
plement of one female. 

June 16, 1930, San Benito, Tex. Among the Hypopachtis cuneits disgorged 
from Thamnophis sauritus proximus were several transformed Bujo valliceps, 
9, 10, 10, 10, 10.5, n, ii, 11.5, 12 mm. Around edges of pond were many more 
but most were away from edges as were some larger ones (16, 19, 19 mm.). 

Journal notes: April 2, 1925, Cotulla Free Camp Ground. Here in an over- 



TOADS: BUFONIDAE 



205 




Plate XL//. Bufo vallicefs. 1,2,4- Males 
(X%). 3- Male croaking (X%) 



XLII1. Bufo tvoodhousii wood- 
housii (X%)- i. 2 - Females ' 34 Males ' 



206 HANDBOOK OF FROGS AND TOADS 

flow cove of Nueces River where water purslane grew were several male Bufo 
valliceps. Saw a female near the shore but was going away when we saw her. 
Possibly our light was responsible. Some 10 or 15 males were calling vigor- 
ously. 

April 22, San Benito, Tex. Pond in a mesquite region. On the east side of 
this beautiful blue water-lily pond we found B. valliceps on the moist earth, 
transforming. Soon after transformation the toads show the light line of tu- 
bercles on the side and the white spot below the eye. They do not show the 
furrowed mterorbital at this small size. 

May 6, Helotes, Tex. Tonight at 9 P.M., as we approached the pond, we 
espied a small head of Thamnophis proxima. A few moments later two larger 
water snakes were close together. They were after a near-by croaking toad. A 
little farther on we heard two male B. valliceps. Presently we saw something 
rolling over and over in the water. It was a water snake. In the semidarkness 
I scooped up the snake and all. The snake dropped a toad. The toad hopped 
limply away. Farther on we found a young Natrix rhombifera beside another 
pair. 

June 16-20, 1930, Brownsville. All along the river in its high state heard 
B. valliceps at night. It is the amphibian note of the river, as is S. campi of the 
dooryards. 

Authorities' corner: 
P. H. Pope, Tex., 1919, pp. 94-95. 

Rocky Mountain Toad, Woodhouse's Toad, American Toad 

Bttfo woodhousii woodhousii Girard. Plate XLIII; Map 14. 

Range: Southeastern Oregon through Idaho, Montana, South Dakota, to 
western Iowa and western Missouri, south through Kansas, Oklahoma, and 
Texas to Mexico, west to Imperial Valley and up Colorado Valley to south- 
east Nevada and through Utah to Idaho. 

Habitat: This toad lives in canyons in mountains and on plains along irri- 
gating ditches. It is also found along rivers and in swamps. In fact its habitat 
is very diverse, being any place where sufficient moisture obtains. 

"The habitat of Bitjo woodhousii is by no means restricted to mountainous 
regions, but includes surroundings as diverse as the sagebrush flats of eastern 
Montana, the prairie fields among the chalk cliffs of western Kansas, the 
Hudsonian Zone mountain sides of eastern Colorado, the irrigation ditches 
that traverse the mesquite plains of New Mexico, and the bottomlands along 
the Colorado River near Yuma, Arizona. During May and June, according 
to locality, adults of this species may be found breeding in shallow sluggish 
creeks, in irrigation ditches, or in freshwater pools in the canyons" (R. Kel- 
logg, Gen., 1932, p. 74). 

Size: Adults, 2^-4% inches. Males, 56-99 mm. Females, 58.5-118 mm. 



208 



HANDBOOK OF FROGS AND TOADS 



General appearance: This toad looks much like our eastern Bufo a. ameri- 
canus but is larger. The general color is grayish drab on the back, with several 
large brownish warts that are usually surrounded by a slightly larger blackish 
area. There is a narrow, light middorsal stripe. The sides are marked with 
several black spots. The crests on the head are dark in color, but not particu- 
larly conspicuous. The tubercles and tips of fingers and toes are reddish 
brown. The head is short and thick. 

H. M. Smith (Kans., 1934, p. 450), who has given us our best account of 
this species, contrasts this form with Bufo a. americantts. 



a. atnertcantis 

1. Warts on body larger, less nu- 
merous. 

2. Usually but one or two warts to 
a dark color spot. 



3. Skin on median anterior surface 
of tarsi and metatarsi with black- 
ish spines. 

4. Parotoids broad and closely ap- 
proximated, not separated by 
more than their own length. 

5. Cranial crests never swollen to 
form a "plateau." 

6. Snouts of males in lateral profile 
pointed to some extent. 

7. Belly usually profusely spotted. 

8. Song a high trill of long duration 
20-30 seconds or so. 

9. Eggs laid single file, enclosed in 
a double tubular membrane, with 
a partition between each egg. 

10. A median dorsal light line rarely 
present; when present very irreg- 
ular. 

n. Parotoids usually separated from 
postorbital ridge; the latter, either 
directly or by a secondary arm, in 
contact with the tympanum. 

12. Second subarticular tubercle of 
fourth toe frequently divided; 
first but seldom not divided. 



w. woodhoiisii 

1. Warts on body smaller, more nu- 
merous. 

2. Some of dorsal spots including 
many warts (eastern specimens), 
or but one or two (western speci- 
mens). 

3. Skin on median anterior surface 
of tarsi and metatarsi without 
blackish spines. 

4. Parotoids narrow, although not 
so long, and separated frequently 
by more than their own length. 

5. Cranial crests frequently swollen 
forming a "plateau." 

6. Snouts of both males and females 
sharply truncate in lateral profile. 

7. Belly usually immaculate or with 
a single median dark breast spot. 

8. Song a low trill of 3-4 seconds or 
so. 

9. No partitions between eggs; but 
a single tubular membrane. 

10. A median dorsal light line always 
present. 

11. Parotoids usually in contact with 
the postorbital ridge, the tympa- 
num separated distinctly from the 
latter. 

12. Second subarticular tubercle ap- 
parently never divided; first but 
seldom. 



TOADS: BUFONIDAE 



209 



Color: Female. Ephraim, Utah, Aug. 19, 1929. Back light grayish olive 
becoming on sides and fore and hind legs tea green. Tympanum pea green, 
parotoid vetiver green or light grayish olive. Crests clove brown or fuscous. 
Spots on upper parts are fuscous, becoming on sides dark ivy green or dull 
greenish black. No very regular spots on either side of back. Each fuscous 
spot on dorsum wart centered, the wart being drab or hair brown. On sides 
spotting more pronounced. Forearm with at least two oblique crossbars. Tibia 
with two or three indistinct crossbars. Under parts deep olive-buff or cream- 
buff. Underside of hind legs pinkish cinnamon. Iris fuscous spotted with 
vinaceous-cinnamon or pinkish cinnamon and sulphur yellow or marguerite 
yellow; pupil rim broken behind and in front, marguerite yellow. 

Ephraim, Utah, Sept. 6, 1929. Female. Throat cream color or pale chalced- 
ony yellow becoming warm buff on breast, cinnamon-buff on belly, and clay 
color on lower side of femur and lower belly. Underside of hind legs reed 
yellow. Underside of forelegs like breast. Upper parts mineral gray or smoke 
gray becoming on fore limbs, side of face, and hind legs tea green or light gray- 
ish olive to grayish olive. Spots on sides most prominent with grayish olive or 
vetiver green dots in them. Spots on dorsum small and warts not large. Line 
down back faint, white for front half. Eye black; pupil rim sulphur yellow or 
light vinaceous-fawn, upper eye with prominent band of vinaceous-buff. Inner 
metatarsal black or clove brown. 

"San Francisco Mts. From fresh specimens: Above pale olive-green with 
a somewhat lighter stripe down the middle of the back; tip of tubercles pale 
red surrounded by black rings; lower surface olive- white" (L. Stejneger, 
Ariz., 1890, p. 116). 

Structure: Cranial crests, prominent, forming right angle back of orbit; 
longitudinal ridges almost parallel; supratympanic or preparotoid ridge ab- 
sent; parotoid glands long, slender, divergent; interparotoid space more than 
twice the interorbital space; two metatarsal tubercles, one very large, one very 
small; throat of male black from line of angle of mouth forward; first finger 
slightly longer than second; large warts on back, each with several pits; this 
toad is larger than B. americantts. 

Voice: The vocal sac is a rounded throat pouch. The call is a vibrating note 
of high pitch, sweet and musical. 

Breeding: This toad breeds from March to July. "The eggs . . . very 
closely resemble those of jowleri. The inner membrane is absent, there being 
but a single tube, about 3.5 mm. in diameter. The egg complement of a single 
female which laid in captivity was 25,644 by actual count" (H. M. Smith, 
Kans., 1934, p. 452). They transform March 25 to September at %-% inch 
(10-13.5 mm.). 

On May 19, 1942, at Beaver Dam Lodge, Ariz., we took about 6 adults and 
about 23 transformed and transforming specimens. These last varied from 
13.5-17 mm. in length, with a mean and also average of 15 mm. 



210 HANDBOOK OF FROGS AND TOADS 

Journal notes: July 24, 1925. At Duncan, Ariz., the engineer at the electric 
light plant told me of several "big" toads at the plant. We went over. They 
were all female Bufo woodhousii. He said they would hop up on the doorsill 
and wait for insects to drop from the wall below the light. Why are they all 
females? Have the males gone to ponds since the recent rains? All these fe- 
males are ripe, not spent. 

May 20, Beaver Dam Lodge, Littlefield, Ariz. Went down to the creek. 
Everywhere toad tadpoles of one sort. Never saw so many. Thousands, hi the 
ditches and in the stream countless tadpoles, large for toad tadpoles. At first 
I thought I had Rana or Scaphtopus tads. Many of these tads were transform- 
ing or transformed. They would take to the stream, and the toadlets had a 
hard time stemming the current. We would go along the bank and cause 
them to jump in, and then we scooped them with the net. The tadpoles were 
in the clear drainage ditches, in the big pond in the creek. Here on the algae, 
not quite emergent, the tadpoles assembled in great numbers. On the algae 
mats that were solid the toadlets could leap along in fine shape. Went out in 
fields next to the creek and caught ten Bufo woodhousii. Tonight at 9 o'clock 
went to the bridge and could hear plenty of Bufo woodhousii. It is a light trill, 
not feeble but sweet, of the quality of the Bufo amcricanus call. They are not 
in chorus. Individual calls not far-carrying. In the closed bag the note of 
woodhousii is a friendly chuckle. 

July 2. Went up Ramsay Canyon, Ariz. Went to Newmans. Here we saw 
B. woodhousii leaping along in the bright sun of midday. It was a scrawny 
individual. 

July 16, below Albuquerque. Last night at camp a Bufo woodhousii feeding 
around our car. 

Authorities' corner: 

W. P. Taylor, Nev., 1912, pp. 344-345. G. A. Moore and C. C. Rigney, Okla., 
J. M. Linsdale, Kans., 1927, pp. 75- 1942, p. 78. 

76. J. C. Marr, Gen., 1944, p. 480. 

Fowler's Toad, Danver's Toad 

Bufo woodhousii jowlcri (Hinckley). Plate XLIV; Map 14. 

Range: New Hampshire to eastern Texas, eastern Arkansas, Missouri, 
southeastern Iowa, eastward in Michigan through Ohio, West Virginia to 
coast. Extensions up Hudson, Delaware, Susquehanna, Ohio, and other rivers. 
Many such Pennsylvania and New York records. Often where B. a. amcri- 
canus and B. w. jowlcri are co-existent B. w. jowlcri is of the river, stream 
banks, or lake beaches; B. a. americanus of the hilly or mountainous regions 
near by. Except for the Northeast our range closely accords with Blair's range. 
Intermediates or misidentifications with B. a. americanus, B. terrestris, and 
B. w. woodhousii make its range difficult to determine at this date. 



TOADS: BUFONIDAE 211 

Habitat: Beaches, coasts, lake shores, or river banks, which are the more 
sandy and warmer places throughout its range, are the usual choice. It is 
common along roadsides, about homes, in fields, pastures, and gardens, in 
sand dunes and pine barrens. It breeds in the shallow water of permanent 
ponds, in flooded low ground or roadside ditches, or along river shores. 

Size: Adults, 2-3/4 inches. Males, 51-74.5 mm. Females, 56-82 mm. 

General appearance: These toads generally have a greenish cast with a yel- 
lowish or buff middorsal stripe. The back is marked by distinct black-edged 
dark spots. In a pair, the male Bufo jowleri is usually darker than the female. 
Live toads sent us from Virginia we described as follows: These are small 
toads with low crests. One is dull greenish in color and one reddish brown, 
each with a light middorsal line. The warts are small and rounded, with sev- 
eral grouped in each dark spot of the dorsal pattern. The parotoids are elliptic 
and nearest together at their midpoints. The ventral surfaces are buffy. The 
throats of the male are greenish black. One toad has the black pectoral spot, 
another one lacks it. Both toads have dark bands along the sides. One has 
much yellow in groin, on rear of the femur, and on tibia and tarsus. 

Color: Lakewood, N.J., from W. H. Caulwell, May 9, 1930. Male. General 
appearance "dark greenish." Dorsal color citrine drab, parotoids buffy brown, 
drab, or wood brown. Top of hind legs and forelegs like dorsum or deep 
olive. Rear of femur, lower half, dark olive or black. Rear of unexposed femur 
with pinard yellow or straw yellow. Spots larger and most conspicuous on 
rear edge of tibia between bars. Groin with some barium yellow or deeper 
yellow spots. Oblique bar on upper eyelids meeting or not on meson. Spot 
just inside of forward end of parotoid, pair of spots midway near meson, 
larger pair near rear end. Last two pairs sometimes united on either side. Pair 
of big spots near meson in middle of back. Other pairs frequently present. 
These spots, two bars on tibia and one on femur, and bars on arm are olive or 
dark olive. Several warts to each spot drab or light grayish olive. Dark spot 
on lower rear of parotoid connected or not with irregular lateral band of dark 
grayish olive or olive. Oblique dark spot from over tympanum to arm inser- 
tion. Side spotted or with dark lateral band mentioned above. Area above 
band and dorsal streaks cartridge buff, ecru-olive, olive-buff, cream-buff, or 
picric yellow. Eye to angle of jaw spot like dorsal ones. Under parts white. 
Pectoral spot of olive or black. Throat deep grayish olive or dark grayish olive. 
Iris rim reed yellow or straw yellow; iris black spotted with light ochraceous- 
salmon, or rufous. 

Female. Dorsal color light grayish olive. Parotoids drab. Bars on eyelids 
and dorsal spots olivaceous black (i) with warts fawn color, army brown, 
or orange-cinnamon ("reddish"). Dorsal streak cartridge buff or pale pinkish 
buff. Light area along side cream-buff or cartridge buff. Warts on hind legs 
reddish in this individual. Belly cream-buff. Black pectoral spot. Throat like 
rest of under parts. 



212 HANDBOOK OF FROGS AND TOADS 

Structure: Crests variable, at times forming a boss; adults never reaching 
the greater size of B. a. americanus; warts on back small and uniform; no pre- 
parotoid ridge from parotoid to postorbital. 

Voice: "The usual note of Fowler's toad is a brief, penetrating, droning 
scream. Only once have I heard a decided departure from this. I heard this 
note late in April in Gwinnett Co., in upper Georgia. A single individual of 
a noisy congregation of males had the unmistakable trill of the common toad, 
but short and decisive like the Fowler's song. It was a perfect combination of 
the notes of both" (H. A. Allard, Ga., 1908, pp. 655-656). 

"While we do not agree with Mr. Allard in calling the song of Fowler's 
toad a 'scream' or 'wail/ it certainly has much less music to it than the trill of 
the American toad. The notes are more closely connected, so that a sort of 
buzzing is produced" (W. DeW. Miller and J. Chapin, N.J., 1910, p. 



"A male toad is a persistent singer during its stay in the water. Its song is 
a combination of a low whistle and a moan, and the two sounds do not melt 
into a chord. The combined sound is discordant and decidedly unpleasant 
to a musical ear, but at a distance the sound is more pleasant, for the moan 
is not apparent and only the whistle is heard. The sound lasts from two to 
three seconds, and may be repeated at intervals of about ten seconds. In 1911 
many sang in the daytime, but in 1912 and 1913 very few were heard except at 
night" (F. Overton, N.Y., 1914, p. 27). 

"One of the most striking differences between the two species lies in their 
voices. While that of the Common Toad is high-pitched and musical, the note 
of Fowler's Toad is nasal, and lower in pitch. Like the voice of the Common 
Toad, it carries well and may be heard at a considerable distance" (K. P. 
Schmidt, 111., 1929, p. 9). 

Breeding: This form breeds from April 15 or earlier to mid-August. In a 
given locality it breeds later than the American toad. The eggs are in long 
files, crowded at first in a double row, and numbering as many as 8000. The 
e gg is %r>-M o inch (1.0-1.4 mm -)> the outer tube 1 /io-'/'i inch (2.6-4.6 mm.), 
the inner tube absent. The tadpole is small, \V\ 2 inches (27 mm.), its greatest 
width toward the rear of the body. The tail crests are low, the tooth ridges %. 
After 40 to 60 days, the tadpoles transform from mid-June to August or later 
at %-% inch (7.5-11.5 mm.). 

"The egg strings, which resemble those of the American toad except that 
the gelatinous tube shows no distinct inner layer nor partitions separating the 
eggs, have been noted in Ohio as early as May and as late as June 24" (C. F. 
Walker, Ohio, 1946, p. 35). 

Notes by S. C. Bishop and Walter Schoonmaker: "The eggs of the Fowler's 
toad, Bujo jowleri (Carman), are laid later than the American toad, Bufo 
americanus (Holbrook). We collected some of the eggs of the Fowler's toad 
on the i3th of May, 1924, at Raft's Pond, near Albany, New York. From these 



TOADS: BUFONIDAfi 

we made the following observations. The eggs of the Fowler's toad, unlike 
the American toad, are sometimes laid in the double string. 

"May 14. The eggs were segmented. Later they became elongated. 

"May 15. Hatched. The tadpoles were hanging on by their suckerhke mouth 
parts, tails down. 

"May 1 6. Grown considerably larger. 

"May 17. Length 7 mm. Gills present. 

"May 18, 11:30 A.M. Actual length 8.5 mm. Gills larger. 

"May 19, 9:55 A.M. Length 10.5 mm. Gills gone from both sides. Mouth parts 
developed. 

"May 20. Length 11.5 mm, 

"May 21. Length 12.5 mm. 

"May 26. Length 14 mm. 

"June 12. Length 20.5 mm. Hind legs present. 

"June 16. Length 21 mm. Hind legs very well developed. 

"June 20. Length 21 mm. 

"June 24. Length 21 mm. Hind legs well developed. Front legs also devel- 
oped. Shape of body changed greatly, also smaller. Tail beginning to shorten. 

"June 28. Tail nearly gone m some specimens and in others the tail is en- 
tirely gone. Some specimens grow faster than others. The young now look 
like the adult. 

"On May 19, 1924, at the pond more eggs were laid, embryos elongated May 
2ist, and hatching May 22nd, and July 1-2, little toads leaving ponds. Males 
began calling May 13, heard May 19, noted calling at intervals until July 2." 

The pair brought to the authors were near the maximum sizes for the spe- 
cies, the male being 70 mm. and the female 8n mm. in body length. 

Journal notes: June i, 1917. About 6 miles beyond Dmwiddie, Va., near the 
road, found several files of toad's eggs. In one case the string was strung out 
in a file 8 or 10 feet long in the current. In another case, in a backwater, the 
mass was tangled around sticks. In most cases the file seems to contain a 
double row of eggs. The note we questioned yesterday evening was Bufo 
jowleri. It is quite different from the sweet droning note of Bttfo americanus. 
Went out at night with a flashlight and captured many B. jowleri and a few 
mated pairs. One pair laid in the fish can overnight. The embrace is axillary. 

April 15, 1921. Air temperature 68F. Tonight at 9 o'clock, Drs. Vernon 
and Julia Haber took me out to the Oakwood Cemetery, East Raleigh, N.C., 
and to St. Augustine grounds. Just beyond, we heard a chorus of what I at 
first mistook for the "bleat" of Microhyla carohnensis. It surely is not the 
note of our northern Bufo americanus, which is a sweeter drone than that of 
B. jowleri. In shallow water along a little drain were plenty of them. The call 
is a striking nasal whir-r-r-r. The Habers say "like a lamb." 

April 15, Raleigh, N.C. Fowler's toads were still in chorus and strongly 
breeding, though started much earlier. The male of the mated pair has the 



214 



HANDBOOK OF FROGS AND TOADS 



first two fingers doubled back and dug into the axils of the female. Often the 
other two fingers may not be doubled back but lie next to the belly of the 
female, or sometimes these two fingers will rest on the shoulder insertion with 
only the first two in the axil proper. The pairs brought in were not laying at 
3 A.M. but at 6:45 A.M. they were well along in oviposition. Water temperature 
at which they were laying is 67 F. 

April 16, St. Augustine grounds, Raleigh, N.C. In the stream near the edge 
found Fowler's toad eggs wrapped around plants. They were in shallow tur- 
bid areas, cattle-punched, and in water 2 to 4 inches deep; many were laid 
last night. They were among speedwell, chickweed, smartweed, marshy St.- 
John's-wort, and other plants, hardly any of which were more than 3 or 4 
inches high at this season. 

June u, 1930, Tickfaw River country, La. At night found Bufo valltceps 
and Bufo jowlen in same woods. Found and heard B. fowlen in ditches. This 
form is Bufo jowlen in voice and structural characters; in some ways not 
typical in color. Viosca says that Bufo valltceps and Bufo fowlen may possibly 
interbreed in Louisiana. 

Authorities' corner: Much printer's ink has been employed on the differ- 
ences between B. w. jowlen and Bufo a. amertcanus. No one has known the 
form more intimately than W. DeWitt Miller. Two authors who helped 
clarify the differential characters are G. S. Myers (Ind., 1927, pp. 50-53) and 
M. G. Netting (Gen., 1930, pp. 437-443). At the same time, 1927-1930, we in- 
dependently assembled these characters in manuscript (15 pp.). We regret 
that space necessitates the elimination of this material, but we include refer- 
ences on the points of discussion. 

READING REFERENCES TO 1930 



1. Voice 

a. S. Garman, Check List, 1884, p. 
42. 

b. F. Overton, N.Y., 1914, p. 27. 

c. M. Brady, Va., 1927, p. 27. 

d. Almost every author on the 
species. 

2. Crests 

a. S. Garman, C.L., 1884, p. 42. 

b. E. D. Cope, Gen., 1889, pp. 278, 
279. 

c. M. C. Dickerson, Gen., 1906, p. 

97- 

d. R. F. Deckert, Gen., 1917, p. 114. 

e. E. R. Dunn, N.C., 1917, p. 621. 

f. M. G. Netting, Gen., 1930, p. 

439- 



Size 

a. S. Garman, sec above. 

b. O. P. Hay, Ind., 1892, p. 458. 

c. W. DeW. Miller and J. Chapin, 
N.J., 1910, p. 458. 

d. E. R. Dunn, N.C., 1917, p. 621. * 

e. M. Brady, Va., 1927, p. 27. 
Head 

a. E. D. Cope, Gen., 1889, p. 278. 

b. A. G. Ruthven, Mich., 1917, p. 4. 
Length of leg 

a. O. P. Hay, Ind., 1892, pp. 458- 

459- 

b. R. F. Deckert, Gen., 1917, p. 114. 

c. A. G. Ruthven, see above. 

d. W. DeW. Miller and J. Chapin, 
N.J., 1910, p. 316. 



TOADS: BUFONIDAE 



215 



e. M. G. Netting, Gen., 1930, p. 438. 

6. Color 

a. O. P. Hay, Ind., 1892, p. 458. 

b. M. C. Dickerson, Gen., 1906, p. 
96. 

c. W. DeW. Miller and J. Chapin, 
N.J., 1910, p. 316. 

d. R. F. Deckert, Gen., 1917, pp. 
113,114. 

e. E. B. S. Logier, Ont., 1925, p. 94. 

f. G. I. Myers, Ind., 1927!), pp. 51, 

5*. 

g. K. P. Schmidt, 111., 1929, pp. 8-9. 

h. Many more writers. 

7. Vertebral line 

a. O. P. Hay, Ind., 1892, p. 458. 

b. M. G. Netting, 1930, p. 441. 

8. Dorsal pattern 

a. M. C. Dickerson, Gen., 1906, p. 

9 6. 

b. G. S. Myers, Ind., 1927^ pp. 51- 

52- 

9. Under parts 

a. M. C. Dickerson, Gen., 1906, p. 
96. 

b. H. A. Allard, Ga., 1907, p. 381. 

c. W. DeW. Miller and J, Chapin, 
N.J., 1910, p. 316. 

d. E. R. Dunn, N.C., 1917, P- 622. 

e. G. S. Myers, Ind., 1927^ p. 52. 

f. M. G. Netting, Gen., 1930, p. 441. 
10. Iris 

a. W. DeW. Miller and J. Chapin, 
N.J., 1910, p. 316. 

b. A. G. Ruthven, Mich., 1917, p. 4. 

c. G. S. Myers, Ind., 1927^ pp. 51- 
52. 

n. Warts 

a. M. C. Dickerson, Gen., 1906, p. 

97- 




Plate XLIV. Bufo woodhousii fowlerL 



b. H. A. Allard, Ga., 1907, p. 304. 

c. W. DeW. Miller and J. Chapin, ^,5- Males (X%). 2. Adult (Xtt). 3- 
XT T * remale 

N.J., 1910, p. 317- 

d. E. R. Dunn, N.C., 1917, p. 621. 

e. R. F. Deckert, Gen., 1917, p. 114- 

f. G. S. Myers, Ind., 1927^ p. 51. 

g. K. P. Schmidt, 111., 1929, p. 9. 



2l6 



HANDBOOK OF FROGS AND TOADS 



h. M. G. Netting, Gen., 1930, p. 17. Eggs 



440. 
12. Muzzle 

a. M. C. Dickerson, Gen., 1906, p. 



1 8. Palettes 

a. M. G. Netting, Gen., 1930, pp. 



439> 44- 
96. 19. Range 

b. R. F. Deckert, Gen., 1917, p. 114. a. K. P. Schmidt, La., 1920, p. 85. 

c. E. R. Dunn, N.C., 1917, p. 621. b. G. S. Myers, Ind., 1927^ pp. 51, 

52. 
c. Many more writers. 



d. M. G. Netting, Gen., 1930, p. 

440. 
13. Parotoids 20. Bars on hind legs 

a. M. C. Dickerson, Gen., 1906, pp. a. M. G. Netting, Gen., 1930, p. 441. 
96, 97. 21. Supra-anal warts 

b. A. G. Ruthven, Mich., 1917, p. i. a. M. G. Netting, Gen., 1930, p. 441. 

c. E. R. Dunn, N.C., 1917, p. 621. 22. Hybridization 

a. W. DcW. Miller and J. Chapin, 
N.J., 1910, p. 317. 

b. C. L. Hubbs, 111., 1918,^43. 

c. A. L. Pickens, S.C., 19273, pp. 



d. K. P. Schmidt, 111., 1929, p. 9. 

14. Shape 

a. H. A. Allard, Ga., 1907, p. 384. 

15. Odor 

a. W. DeW. Miller and J. Chapin, 
N.J., 1910, p. 316. 

1 6. Breeding season 

a. All authors agree the breeding 
season is 2-4 weeks later than in 



25, 26. 

d. A. L. Pickens, S.C., 1927^ p. 
109. 

e. M. G. Netting, Gen., 1930, p. 
442. 



Bufo a. amcricanus. 

For five recent papers on B. w. jowleri, see A. P. Blair (Gen., i943a), M. G. 
Netting and C. J. Coin (Fla., 1945), C. F. Walker (Ohio, 1946), R. A. Littleford 
(Md. 1946), and A. P. Blair (N.J., 1947)- 



FAMILY HYLIDAE 

Genus ACR1S Dumeril and Bibron 
Map 15 

Cricket Frog, Savannah Cricket, Cricket Hylodes, Peeper, Southeastern 
Cricket Frog, Cricket Hyla, Sphagnum Cricket Frog, Coastal Cricket Frog 

Acris gryttus gryllus (Le Conte). Plate XLV; Map 15. 

Range: Dunn (Gen., 1939, PP- 1 53~ 1 54) g^ ves southeastern Virginia (Nor- 
folk) to Florida to parishes of Louisiana. 

Habitat: Terrestrial, shade-loving frog. In meadows or about creeks or 
ponds in the open vegetation mats or wooded edges. 

"All in the region of coastal sphagnum swamps" (C. S. Brimley, N.C., 
1940, p. 15). 

"The most abundant amphibian of the region and the only one which 
could be taken during every month of the year. Each roadside pool or ditch 
contained numerous individuals and the swamps and marshes literally 
swarmed with them. In the short winter season, here lasting from about the 
middle of December till the first week or two in February, Cricket Frogs 
were much less in evidence than at other times, and as collecting is then at its 
easiest it is clear that the majority hibernate. But even on the coldest winter 
days no difficulty attends the capture of a plentiful supply, which thus pro- 
vides a dependable source of food for many of the snakes kept in laboratory 
cages throughout the year. Sporadically met with in the piedmont" (J. D. 
Corrington, S.C., 1929, p. 65). 

"Almost any aquatic situation; commonest in marshes and ditches with 
shallow margins choked with hydrophytes" (A. F. Carr, Jr., Fla., i94ob, 

P-55)- 

Size: Adults, %-i% inches. Males, 15-29 mm. Females, 16-33 mm - 
General appearance: This is a small tree frog, but it looks like a small true 
frog (Rana). It varies in color, black, dark brown, reddish brown, light 
brown, green, or gray; or the markings may be reddish on a green ground. 
Between the eyes there is usually a dark triangle, white-bordered behind. This 
species possesses rear femoral stripes, oblique bars on the sides, light spots on 
the jaw, and an oblique white stripe from the eye to the arm. The skin is 
more or less tubercular. 
"Much like the preceding [A. g. crepitans}, but the head is longer and 



2l8 



HANDBOOK OF FROGS AND TOADS 



more pointed, the dark triangle between the eyes is longer and more acute 
behind; the edge of the upper jaw is dark with three or four light vertical bars 
or spots on each side and the dark stripe on the back of the thigh is darker 
and more constant. Slightly smaller and slimmer than the common cricket 
frog. This species averages darker and smaller than the preceding, the darker 
color probably owing to the darker soil of the region it inhabits, and I have 
seen but few wholly green specimens. In habits it is similar, being quite as 




Map 75 

active if not more so. Both species occur in both shady and sunny situatioas. 
By Cope this was considered a more Southern race of the preceding, but most 
authors have recognized it as merely an individual variation. Dr. Percy 
Viosca, Jr., however, seems to have shown fairly conclusively that they are 
two distinct species and I agree with him. Dunn (1939) says the difference 
in the amount of the webbing on the hind feet is the best criterion by which to 
separate the two, and I find in looking over some fifty specimens in the State 
Museum, about evenly divided between the two forms, that a glance at the 
first or shortest toe is usually sufficient to distinguish either; if the webbing 
extends to the tip, the frog is crepitans, if the last joint is free from web it is 
gryttus" (C. S. Brimley, N.C., 1940, p. 15). 



TREE FROGS: HYLIDAE 



219 



Color: On one day, April 23, 1921, in Georgia, I saw black, dark brown, 
reddish brown, light brown, green, and gray specimens of Acris. 

Okefinokee Swamp, Ga., June 25, 1921. Male. Stripe down back and around 
triangle dark olive-buff. Throat raw sienna. All over the throat are collections 
of dark dots, sometimes arranged in a reticulate fashion. Iris pale vinaceous- 
drab on black, light orange-yellow pupil rim. 

Female. Clove brown above; triangular spot between eyes obscured by this 
dark color; throat, breast, and belly pale olive-buff; more or less of same color 
on underside of forelegs, the spot below eye, along the upper and lower jaw, 
and the line from eye to arm insertion; area back of arm insertion pallid vma- 
ceous-drab; oblique bar on side clove brown with pale olive-buff and olive- 
ocher; underside of hind legs clear with little pigment; long stripe on rear of 
femur, snuff or Dresden brown with clear unpigmented stripe below and 
above; also another brown stripe above the upper clear area; from vent 
around bases of hind legs to venter are pale olive-buff papillae. 

Structure: Tympanum indistinct; tympanic fold present; fold across breast 
frequently present; disk small; hind limb very long; tibia very long. 

"Width of head across the base of lower jaw less than its length from that 
point to tip of snout, legs longer and hind feet less webbed, the web of the first 
toe absent from the last joint; heel when leg is extended reaching beyond the 
tip of the snout. Sphagnum cricket frog (Acris gryllus)" (C. S. Brimley, N.C., 
1940, p. 15). 

"Webbing on toes less extensive; last three phalanges of fourth toe free from 
webbing; usually with third and fifth toes not reaching middle of the third 
phalanx from tip of fourth toe. Acris g. gryllus" (E. B. Chamberlain, S.C., 
1939, p. 12). 

Voice: Characterizations of the Acris call are numerous. 

April 1 6, 1921, at Raleigh, N.C. Acris gryllus call sounds like a rattle or some 
of the metal clickers, gtcl^ t gtcf(, gicf( f or l^ic^ f(tc^ f fycl(, in rapid succession. 
Frequently one finds the males with inflated vocal sacs even when not calling. 
When calling, the throat is never fully deflated. After a call it may be swollen 
to three-quarters its full capacity. Then when the call is given, the body sides 
are compressed and the vocal sac is extended to its limit. 

Holbrook (Gen., 1842, IV, 132) wrote, "This is a merry little frog, constantly 
chirping like a cricket, even in confinement. . . ." Of his captives he said, 
"Their chirp, at times, was incessant and sprinkling them with water never 
failed to render them more lively and noisy." 

"Call an irregular series of iiks; comparable to sound made by scratching 
teeth of a comb; somewhat cricket-like; ik-ik, ii-ik, ii-ik" (A. F. Carr, Jr., Fla., 
1934, p. 21). 

Breeding: This species breeds from February to October. The single eggs are 
few (250), are brownish and white, and are attached to stems of grass in shal- 
low water or are strewn on the bottom. The egg, %5 inch (0.9-1.0 mm.), has 



220 HANDBOOK OF FROGS AND TOADS 

a single envelope Yio-Vs inch (2.4-3.6 mm.). The dark olive tadpole is me- 
dium, i 1 Me inches (42 mm.), full and deep-bodied, its tail long with acumi- 
nate tip and with a black flagellum. The tooth ridges are %. After 50 to 90 days 
or longer, the tadpoles transform from April to October at %-% inch (9-15 
mm.). 

June i, 1917, at Dinwiddie, Va., we found this species breeding. They had 
chosen a shallow grassy meadow pool i to 4 inches deep. The eggs were at- 
tached singly to sedge stems or were strewn singly on the bottom. In one or 
two cases, three or four eggs were close together. Many of the eggs were in 
water not more than an inch deep. The eggs are firm. We found no more than 
ten eggs. 

"Calling was earliest heard on January 22. This increased to the proportions 
of a chorus by March 17. Clasping was first observed on April 19 while the 
first eggs were procured on April 25" (B. B. Brandt, N.C., 19363, p. 217). 

"They may breed during any month of the year. The eggs are laid in very 
shallow water, often among semiaquatic or terrestrial vegetation which has 
been temporarily inundated" (A. F. Carr, Jr., Fla., 1940, p. 55). 

Journal notes: The Cornell party of Dec. 22, I9i3~jan. i, 1914, found this 
species active and took several specimens. Doubtless this species is more or less 
active throughout the year in the Okefinokee Swamp. 

April 23, 1921, Okefinokee Swamp. Acris captured a lot of them. Some- 
times black on a black soil and hard to see except when they jump. Some brown 
all over back (except for dark marks) when on brown pine needles. Sometimes 
green all over except for the dark marks. Sometimes gray over drier sand. 
Among some of the light brown needles Acris reddish brown even on back 
of fore limbs and hind limbs. 

A few of our notes on the jumping records of Acris follow: 

April 23, 1921. Acris usually jumps for several leaps before it disappears. 
April 25. Acris males jump 3 feet at a time on the water's surface. May 17. Acris 
can leap at least my own pace. May 21. Acris has been hopping around on the 
ground and into small bushes from the ground and down to the ground again. 

Authorities' corner: 

W. Bartram, Gen., 1791, p. 278. C. J. Goin, Fla., 1943, p. 148. 

P. Fountain, Ga., 1901, p. 63. P. Viosca, Jr., La., 1944, p. 55. 

R. F. Deckert, Fla., 19153, p. 22. G. L. Orton, La., 1947, pp. 377-378. 

A. F. Carr, Jr., Fla., 1940, p. 55. 

Cricket Frog, Western Cricket Frog, Valley Cricket Frog, Western Cricket, 
Peeper, Savanna Cricket Frog, Rattler 

Acris gryllus crepitans Baird. Plate XL VI; Map 15. 

Range: Dunn (Gen., 1939, pp. 153-154) gives Connecticut, s< 
New York, New Jersey, Pennsylvania, Maryland, Delaware, northwest to 



TREE FROC&: HYLIDAE 



221 




Plate XLV. Acris gryllu; gryttus (XO- *** ^ LVL Acris 

1,3. Females. 2. Male. 4. Tadpole. (X%)- A5A7- ^males. 3 ,4- Males. 8,9. 

Tadpoles. 



HANDBOOK OF FROGS AND TOADS 

Canada, west to Utah and New Mexico, and south through Virginia to 
Georgia and western Texas. Sea level to 2000 feet. 

Habitat: Swales, lake margins, stream edges, springs, pasture pools. 

"This tiny frog is distributed all over the State wherever there are lakes, 
ponds, springs, or streams. I have found it even in the heart of well populated 
cities in little pools formed by rains. While allied to the true tree frogs, this 
species never climbs trees but lives among water plants and in the vegetation 
along shore. When alarmed it retreats to the water after the manner of a true 
water frog" (J. K. Strecker, Jr., Tex., 1915, p. 49). 

"This is one of the commonest forms found in the Wichita National Forest 
and if an effort were made large numbers could be taken. Several dozen were 
preserved on this trip and some hundreds are in the collections as the result of 
many short collecting trips which have been taken to this forest. They are 
found almost anywhere that water was available and in this region were plenti- 
ful along most parts of the streams and particularly around the edge of Lost 
Lake and other slow-flowing water areas. Their abundance along West Cache 
Creek in spite of numerous campers was very noticeable" (A. I. Ortenburger 
and B. Freeman, Okla., 1930, p. 177). 

"It is generally abundant wherever permanent waters are to be found, where 
it may be taken as early as the middle of February and as late as November" 
(D. A. Boyer and A. A. Heinze, Mo., 1934, p. 189). 

Viosca stated that Acns gryllus abounded along the creeks of his Pine and 
Hardwood Uplands division or "Shortleaf Pine Hills." "Fresh Water Marshes: 
Acris crepitans, although a lowlands species generally, is especially abundant 
here . . ." (P. Viosca, Jr., La., 19233, pp. 36, 39). 

W. S. Blatchley (Ind., 1891, p. 27) calls this species "the most abundant tail- 
less batrachian in the country. Hundreds are to be seen along any small stream 
in spring and autumn. They appear less common in summer." 

"This species was the most abundant amphibian within the limits of this 
area. It was found most abundantly at the edges of bodies of water in all parts 
of the area in spring although some were found on the shores of the lake 
through summer and until late in the fall. On February 2, 1924, several were 
found in the creek above the bridge. They were in the water above some olcl 
ice and below a top layer of new ice. All the frogs were stifl and floating and 
apparently they were dead. In the warm water of the springs and just below 
the springs a few frogs of this species were found throughout the winter" (J. M. 
Linsdale, Kans., 1927, p. 76) . 

Size: We have made no study of this segregated group. A tentative summa- 
tion is: adults, %-i% inches; males, 17 or 18-30 mm.; females, 20-35 mm. Did 
Fowler's (1907) 2% inches mean i/4 inches? 

We measured 20 males and 20 females from Texas and Mississippi Valley 
with these results: males^ 17.5-29.0 mm., average 21 mm., mean 23 mm.; fe- 
males, 19-28 mm., average 24 mm., mean 23 mm. In one lot a ig-mm. male 



TREE FROGS: HYLIDAE 223 

was debatable; in another lot three individual specimens, 17, 18, 18 mm., were 
hard to sex; in another lot we had some males 17.5 and 18.5 mm. 

General appearance: In 1923 Viosca (La., 19233, p. 43) said: "The puzzling 
status of Acns, as far as Louisiana is concerned, has been positively cleared by 
these studies. There are two distinct species in Louisiana, the upland species 
being, tentatively, Acris gryllus, and that of the lowlands, Acris crepitans. 
Wherever their ranges overlap, they are found side by side without inter- 
breeding, each with its characteristic chorus and habits." Dunn (Gen., 1939, 
PP- I 53" I 54) confirmed Viosca's opinion that two distinct species are involved 
and the distinguishing characters are those he mentioned: "'The best char- 
acter for distinguishing the two species is the amount of webbing of the toes, 
crepitans having much more web.' 

"gryllus crepitans 

Smaller Larger 

Less web (5 phalanges of toe More web (2 to i l /> phalanges of toe 

4 free, toe i partly free) 4 free, toe i completely webbed) 

More rugose Smoother 

Anal warts less prominent Anal warts more prominent 

Legs longer, heel beyond snout Legs shorter, heel not to snout 

Thigh more definitely striped Thigh less definitely striped." 

"If this form actually exists, as Viosca (1923, 1931) maintains, it certainly 
does not exist in Kansas. The specimens examined are too uniform in charac- 
ter to permit of more than one species, and the extent of variation is well 
within that of gryllus' (H. M. Smith, Kans., 1934, p. 461). 

"Dunn (Gen., 1939, p. 154) lists six criteria for use in distinguishing crept- 
tans from gryllus. Four of these, (i) larger size, (2) more extensively webbed 
feet, (3) shorter legs, and (4) less definitely striped thighs, are evident in our 
specimens when they are compared with topotypes of gryllus from Riceboro, 
Georgia. The two other diagnostic characters used by Dunn do not hold in 
our specimens; they seem, therefore, to merit further discussion. First he char- 
acterizes gryllus as 'more rugose' than crepitans; the contrary is true in the 
Rockingham County specimens which are definitely more rugose than are 
gryllus topotypes. Wide variation in the amount of rugosity occurs in both 
species and we feel that this character is the least useful of those used by 
Dunn. Secondly, Dunn states that the anal warts of gryllus are less prominent 
than those of crepitans. This statement is ambiguous because it fails to indi- 
cate whether the warts are prominent by reason of their color, size, or number. 
The startlingly white color of the subanal warts of some specimens of Acris 
is a result of preservation in formalin, as the senior author has determined ex- 
perimentally; specimens preserved in alcohol show less color change and the 
subanal warts do not fade to an ivory white. It is quite evident however that 
rugosity of the central thigh area is more characteristic of crepitans than of 



224 HANDBOOK OF FROGS AND TOADS 

gryllus; the Virginia specimens and many examples of crepitans from else- 
where exhibit a greater number of subanal warts than do topotypes of gryllus, 
but brilliant white warts are present in some specimens of each species. In 
addition to the characters mentioned by Dunn, crepitans has a shorter head 
and a more obtuse snout than gryllus" (M. G. Netting and L. W. Wilson, 
W,Va., 1940, p. 6). 

Color: "Color above, some shade of gray, brown, or olive-green, often with a 
median longitudinal diffuse band of red or green, and with several black spots, 
of which a triangular one between the eyes is constant and characteristic. Be- 
neath pale. Upper jaw black or dark brown, with four vertical pale lines on 
each side. A narrow pale line extends from the lower posterior part of the eye 
to the base of the fore leg. Above this line lies an elongate black spot which ex- 
tends from the eye towards, but does not quite reach, the fore leg. Behind the 
insertion of the fore leg, on the side, is a large oblique black spot margined with 
white. Another similar but smaller spot lies in advance of, and above, the in- 
sertion of the hind leg. The triangular spot between the eyes is narrowly mar- 
gined with white, its apex pointing backward. The middle of the back is often 
occupied by a longitudinal red or green band, and immediately on each side of 
the latter are several obscure black spots. Color beneath pale, sometimes tinged 
with yellow on the throat. Throat more or less speckled with dusky or brown. 
Lower jaw pale, or with a few dark specks at the symphysis, becoming darker 
towards the angle of the mouth, from which point a dark dash passes to and 
upon the base of the fore leg. Legs and digits dark above, with round dark 
spots; pale and unmarked below. A black spot may often be visible over the 
vent, and generally a dark bar passes from this region along the posterior sur- 
face of the thigh" (H. Carman, 111., 1892, p. 341). 

"Acris gryllus. Cricket frogs are common along the shores of all bodies of 
water. Some of them were very large. All were dark in color and most of them 
showed either a red or green streak down the center of the back" (K. P. 
Schmidt, III., 1923, p. 49) . 

Structure: (See A. gryllus gryllus for differences in the two forms.) 

"Width of head across base of lower jaw about equal to distance from that 
point forward to tip of snout; legs shorter and hind feet more fully webbeH, 
the web on the first (shortest) toe reaching its tip; heel when leg is extended 
not reaching tip of snout Common Cricket" (C. S. Brimley, N.C., 1940, 
p. 15). 

"Webbing on toes more extensive; last two phalanges of fourth toe free from 
webbing; usually with third and fifth toes reaching beyond middle of the third 
phalanx from tip of fourth toe. A. g. crepitans" (E. B. Chamberlain, S.C., 1939, 
p. 12). 

"None of this questionable species was found in Beaufort County although 
Brimley (1926) gives the range in North Carolina as 'central and part of east- 
ern districts.' The writer has seen individuals commonly in Durham County 



TREE FROGS: HYLIDAE 225 

which are referable to this species. Viosca (1923) has taken considerable inter- 
est in the genus, as a result of field observations maintaining that in Louisiana 
there are two distinct species of Acris. The writer is inclined to the view that 
more field and experimental study is needed to solve the problem. It was ob- 
served that in Chase's Lake, Brooksville, Mississippi, only typical gryllus forms 
occur while at Cryme's Pond both forms occur with calls which are recogniza- 
bly different. The former locality is in a Lafayette Red Clay area while the 
latter, less than four miles distant, is in a Selma Chalk area" (B. B. Brandt, 
N.C., 19363, p. 217). 

Voice: Feb. 24, 1925, Helotes, Tex. Tonight air 64 at 8:50 P.M. Heard at 
"Ornate Fork" quite a chorus of Acris, gicl(, gic^ gicf( or l(icl(, %icl{, ^icl{. 

March 8-12, Helotes, Tex. Evenings in great chorus for several days. Creek 
is filled with them. 

"Besides their rattling call a squeaky sound was heard occasionally, though 
only during the breeding season. . . . 

"It may be stated that in my experience their call appears to be variable. The 
usual note is not heard at a great distance, and is described by one writer as 
exactly imitated by striking two marbles together, first slowly, then faster and 
faster, for a succession of about 20 to 30 beats. Perhaps the rattling of castanets 
would be a better suggestion" (H. W. Fowler, N.J., 1907, pp. 102-103). 

"The name Cricket Frog was given to it, on account of its song, which bears 
a strong resemblance to the chirping of the black cricket. These tiny frogs sing 
in chorus in spring. The sound can be imitated by striking together two peb- 
bles or two marbles, beginning slowly and continuing more rapidly for thirty 
or forty strokes. The male frog is the singer and in doing so inflates his yellow 
throat enormously. . . . The first warm days in early spring bring them out. 
Feb. 14, 20; Mar. 5; May i; Sept. 7; Oct. 16" (J. Hurter, Mo., 1911, p. 
102). 

"Feb. 8, 1918. Buffalo Bayou. They are fairly abundant all along the banks 
and are calling vigorously. The call is a soft trill resembling the tree cricket or 
the mole cricket" (P. H. Pope, Tex., 1919, p. 97). 

"The cricket-like chirping of these abundant little frogs is a familiar sound 
about the marshy borders of streams and ponds from April to July. The singer 
is often hard to locate due to the carrying capacity of the sound and a more or 
less ventriloqual effect" (G. E. Hudson, Neb., 1942, p. 25). 

"The voice of a cricket frog is a combination of a rattle and a musical clink, 
but it is only about half as loud as that of a spring peeper. A chorus heard at a 
distance sounds like the jingling of small sleigh bells, for the musical element 
of its call travels farther than the rattle. A chorus heard close by sounds like the 
rattle of small pebbles poured upon a cement pavement. 

"An individual frog sings for from thirty to forty-five seconds at a time. Its 
call has three phases. The first phase lasts for about five seconds and sounds like 
the clicks of a boy's marble dropped upon a cement pavement, once or twice a 



226 HANDBOOK OF FROGS AND TOADS 

second from a height of about six inches. The second phase sounds like the gal- 
loping of a small pony on a brick pavement, or like the clicks of a boy's marble 
dropped upon a pavement from a height of only an inch or two, and allowed 
to bounce each time. The third phase sounds like the regular cree-cree-creeing 
of a tree cricket, or like the rattle of a boy's marble that bounces rapidly when 
it is dropped at frequent intervals from a height of only half an inch. The time 
and rhythm of the sounds are about the same as that of the following syllables 
pronounced with the speed of ordinary reading: click, click, click, click 
. . . click-e-ty, click-e-ty, click-e-ty, click-e-ty, click-e-ty . . . cree, cree, cree, 
cree. . . . 

"The cricket frog inflates a vocal sac under its chin during its call. It often 
sits quietly with its sac distended for many minutes between its calls. The vio- 
lent efforts of its body in producing its sound make the frog resemble a small 
boy on his hands and knees blowing a fire with all his might. The vocal sac is 
bright yellow, and when it is seen distended in the day time, it is so conspicuous 
that it reveals many a singer that otherwise would be almost invisible on a lily 
pad" (F. Overton, N.Y., 1914, p. 31). 

Breeding: Early April-July. (See A. g. gryllus.) H. M. Smith (Kans., 1934, 
p. 389) gives the egg as having a single envelope 2.3 mm. or more. Dr. Kather- 
ine Van Winkle (Mrs. E. L.) Palmer who worked over our Acns collections 
and our 1917 transcontinental material found that in an Ames, Iowa, congress 
of Acris June 12, 1926, some eggs tended to be in bunches or masses. We meas- 
ured her Ames material and found males 21-29 mrn - m length and females 27- 
29 mm. Most of the records indicate transformation from 13-15 mm., but we 
were quizzical and, on testing out our Texas material, found they ran mainly 
from 1T-I2 mm., sometimes 10 or 10.5 and 12.5 or 13 mm. An inclusive range 
of 10-15 mm. is, therefore, better for transformation sizes. In the same way, 
few tadpoles reach 45 or 46 mm.; most run from 30 to 36 mm., with the body of 
the tadpole usually 11.5-14.0 mm. 

Miss Dickerson believed their chorus was loudest in late April and early May 
when they attached their eggs to grass blades or leaves in the water. She stated: 
"The development of this frog is less rapid than that of the Common Tree 
Frog, the Eastern Wood Frog, or the American Toad. The tadpoles may be 
found in the water as late as August. The final transformation takes place in 
September. The young tree frogs, as well as the older ones, seek shelter from 
the cold under stones and leaves at the margins of their brook or marsh." 

"[Eggs are] not deposited in large groups, and not concealed under objects; 
outer envelope usually not over 7.5 cm. Envelope single. Envelope 2.3 mm. or 
more Acris gryllus (Le Conte)" (H. M. Smith, Kans., 1934, p. 389). 

"Although individuals are sometimes collected throughout the winter, and 
are rather abundant in March, their breeding activities do not begin until early 
April. . . . Near Lawrence clasping pairs have been collected as late as May 9 
(1933) ; they probably breed much later, as their songs are frequently heard as 
late as July, and Gloyd (personal notes) has heard them singing about Man- 



TREE FROGS: HYLTDAE 227 

hattan as late as July 15. Eggs have been laid in the laboratory on May 10, from 
females collected the preceding night (1933). The breeding sites chosen are 
varied. In regions where the species is found in abundance, permanent lakes, 
streams and springs always have their quota. Frequently they breed in and sing 
from temporary pools in pastures or at road-sides" (same, p. 459). 

"On July 17, 1932, metamorphosis was occurring in these frogs at Isle Du 
Bois Creek, where, it was estimated, many thousands of these small frogs oc- 
curred, and where specimens in almost all stages of development were present. 
Some juveniles without fully absorbed tails had bright green dorsal patterns, 
while tailed specimens approaching transformation possessed the typical tri- 
angular dorsal marking, but no such larvae were observed to have the black 
tail tip characteristic of the tadpoles occurring in a woodland pond at Danby, 
about two miles distant. A transforming specimen upon emergence from the 
water on August 9 possessed a body length of 14.4 mm., tail 14.9. Two hours 
and forty minutes later the tail measurement was 10.75, an absorption of 4.15 
during this period. Another larva, measured at 9:15 P.M., had a body length of 
15.1, tail 31.7, of which 9 mm. possessed dark pigmentation; the mouth was 
transformed, the dorsal pattern distinct. By 7:30 P.M. August 10 the body 
length was 14.4, tail, 24, dark tip, 3.5 and by 2 P.M. August u the body 
length was 14, tail, 1.5. In this period of 40% hours the reduction in body 
length was i.i mm., tail 30.2. By September 18 the transformation period was 
neanng its end and few larvae could be taken by seining at this time. On this 
date, however, a ribbon snake, Thamnophis sattnttts proxtmns was lying on 
a mat of aquatic vegetation at a pond near Danby, and when captured it dis- 
gorged three cricket frogs, one a transforming specimen" (D. A. Boycr and 
A. A. Hemze, Mo., 1934, P- x< ^9)' 

"The eggs were laid singly or in small masses. One female had 248 . . ." 
(G. E. Hudson, Neb., 1942, p. 25). 

Journal notes: We have seen it from New Jersey to Georgia, Louisiana, 
Texas, and Iowa, but like most workers we have neglected this most ubiqui- 
tous tree frog. Everyone remarks their choruses. We will forbear except to 
give the scanty notes of two seasons in Texas. 

Feb. 8, 1925, Rio Cibola River, Tex. A few Acns called. 

Feb. 8-Feb. 21, Helotes. Once in a while hear a few Acns calling. Not a loud 
full-throated Acrls call yet. 

Feb. 21. Heard them calling, air and water 7oF., 9-10 P.M. 

Feb. 24 to March 8-12. [See "Voice." | 

March 13. In great chorus last night and tonight. In our fork it is very 
common. 

March 16. Last night no Acris calling. 

April i. Heard cricket frogs at Sabinal, at Nueces River, and at Uvalde. 

April 2, Cotulla. Heard in Nueces River or extension many, many Acris. 

May 5, Helotes. On talus slope we turned over a flat stone and saw a greenish 
frog. Another jumped and my eye followed it. It impressed me as Acris. I let 



228 HANDBOOK OF FROGS AND TOADS 

the stone down. Later looked under the stone, and behold a Marnock's frog 
a red-letter day. Later we went to the spot where the other frog jumped and 
it proved to be an Acris. It must have finished breeding to be here. 

May 9, San Antonio, 8:30 P.M. Acris calling strongly near St. Mary's College 
and in Bandera. 

May 14, Boerne. Heard a few Acris. 

May 24, Helotes. Went to Lee's Branch to get Acris crepitans series. Many 
Acris transformed. 

June 6, Devine. Found Acris tadpoles with black-tipped tails. 

Jan. 17, 1942, With Quillms to Classen Ranch. In clear Fork of Cibola . . . 
Roy dug under roots of a tree and brought out a Bufo valliceps. In the creek 
saw plenty of Acris crepitans, tadpoles minute to almost mature of R. pipiens. 
Saw an immense R. catesbeiana male in mass of vegetation. 

July 22, Toyahvale, Tex. Tonight at 9:30 drove to where Cherry Canyon 
goes across the highway and several miles beyond. At first we heard a note 
1 thought was a rattle in rear of our auto, but Anna persisted, and then I got it: 
full Acris crepitans. Got plenty of them for first 2 or 3 miles west of Toyahvale. 

July 23. Trip to Phantom Lake with Alex Izzard of Balmorhca State Park. 
Saw several Acns here and caught them. Water about 70 F. A fine spring. 

August 2. Beyond Carrizo Springs, Tex. At Sycamore Cr. boundary line of 
two counties took one transformed Acris. 

In Texas in 1925 we began trips about February 8, when Acris appeared if 
nothing else. For many entries we merely say, "No end of Acris" "Acris in all 
creeks of whatever size." Always merely "Acris." Not until May 24-26, when 
they were transforming and transformed, did we courageously enter A. crepi- 
tans. At present we are almost willing to grant crepitans and gryllus, but are 
there one to three or more forms of Acris yet undescribed ? This omnipresent 
species may prove more involved than Rana pipiens. 

Authorities 9 corner: 

H. Garman, 111., 1892, p. 342. C. S. Brimley, N.C., 1940, p. 15. 

J. Hurter, Mo., 1911, p. 101. G. A. Moore and C. C. Rigney, Okla., 

T. L. Hankinson, 111., 1917, p. 324. 1942, p. 78. 

B. W. Evermann and H. W. Clark, G. L. Orton, La., 1947, pp. 375-377. 

Ind., 1920, p. 634. 

In a recent paper (May 10, 1947) F. Harper (Gen., 1947) restricts Acris g. 
crepitans to the eastern portion of its credited range, i.e., from Connecticut to 
New York to eastern Texas, and establishes a new form, A. g. blanchardi for 
the cricket frogs from southwestern Michigan to southwestern Missouri to 
southwestern Texas and westward to Utah and Arizona. The new form "is 
distinguished ... by slightly greater linear measurements, by decidedly 
greater bulk, by somewhat more extensive dusky area on the posterior face of 
the femora in the vicinity of the vent." Our preceding discussion of Acris g. 
crepitans is therefore composite if this interpretation proves the correct one. 



Genus PSEUD ACRIS Fitzinger l 
Maps 1 6-1 8 

Mountain Chorus Frog, Chorus Frog, Ohio Chorus Frog 
Pseudacrts brachyphona (Cope). Plate XLVII; Map 16. 

Range: Southwestern Pennsylvania, western Maryland, southeastern Ohio, 
eastern Kentucky, West Virginia, northern Alabama (Viosca), northern Mis- 
sissippi. Must be in Tennessee. 

Habitat: Springy hillsides, grassy pools, ditches, sources, and along upper 
courses of upland rivulets more hilly than lowland habitats. 

Size: Adults, 1-1% inches. Males, 24-52 mm. Females, 27-34 mm - 

General appearance: These arc small frogs, gray or brown in color, medium 
in size for Pseudacns, with the most distinct digital disks of this genus. They 
arc more stocky in body and broader in head than P. n. triseriata and P. n. 
feriarum. The usual middorsal stripe or row of spots is lacking. They often 
have a light middorsal area somewhat after the pattern of the cricket frog 
(Acris gryllus). The intcrorbital triangle is not white-edged behind. The dor- 
solateral bands curve from the eye to midback to groin, making two crescents. 
They often meet in midback to form a cross or transverse bar. Sometimes the 
pattern consists of a cross or a bar alone. 

Color: (See "General appearance," "Structure," and "Authorities' corner.") 
In general color these frogs range from the sorghum brown, deep brownish 
drab, or mars brown of Hyla femoralis to some of the grays found in Acris or 
Hyla femoralis or to the blackish olive of Pseudacns n. triseriata. 

Structure: The original description "A specimen of nearly the si/.e and 
form of Hyla femoralis was taken in western Pennsylvania, near the Kiskimi- 
nitas River. In proportions it does not differ from the Feriarum, but the toes 
arc fringed, the dilations larger and the coloration different. Above blackish- 
ash, abruptly defined on the sides. Lateral band not extending beyond tympa- 
num. No median dorsal band, but two black dorso-laterals of double ordinary 
width converge from each tympanum and extend to end of urostyle inclosing 
with the interorbital triangle a narrow, anteriorly bifurcate dorsal band of 
ground color" (E. D. Cope, Gen., 1889, p. 341). 

Compared to P. ornata, P. n. feriarum, P. n. triseriata, it has a wider head 

1 The two workers who in the last ten to fifteen years have added most to clarification 
of this genus are C. Walker and F. Harper. 



2 3 



HANDBOOK OF FROGS AND TOADS 



and a longer hind limb, tibia, and foot with tarsus. In most ways it falls into 
the Pseudacris ornata group; in some with the Pseudacris nignta group, e.g., 
as shown in the statistics (number of times in length) below: 

P. ornata P. brachyphona P. n. feriarum 
Width 
of head 2.86-3. 2.5-3.1 

0.60-0.67 

i. 82- r. 9 3 



P. n. triseriata 



Hind limb 
Tibia 
Foot with 
tarsus 



2.86-3.4 

0.68-0.75 

2.06-2.28 



3-0-3-33 

0.65-0.74 

2.0-2.26 



3-25-3-5 

0.71-0.75 

2.24-2.54 



1.26-1.42 



1.23-1.60 



I -37- I -57 



These measurements distinguish it from P. n. triseriata, but not so clearly from 
P. n. feriarum. The proportions for P. brachyphona are in three parts very close 



Pseudacris brachyphona 




16 



to those of P. n. nigrita. Although we have put this species in the P. ornata 
group in the key, nevertheless there are some respects in which it is not clearly 
divorced from the P. nigrita six subspecies. 

Voice: "The note of this species is quite different from that of the C. triseria- 
ttts, not being continuous, but in sets of crepitations repeated in time and at in- 
tervals" (E. D. Cope, Gen., 1889, p. 341). When P. n. triseriata and P. bra- 
chyphona are calling they are different; the latter has a faster, higher note, yet 
its call belongs distinctly in quality and form with the P. nigrita group. 



TREE FROGS: HYLIDAE 231 

"The voices of the two (tnsenatits and brachyphona) are much alike but the 
call of brachyphona is given more quickly, with a higher pitch and a different 
quality so that the effect of a chorus is quite distinctive" (C. F. Walker, Ohio, 



"Their notes are repeated at the rate of about 50 to 70 times a minute and 
may be continued for several minutes although they usually stop in fifteen to 
twenty seconds. The call is strong and rapid and on a clear night a chorus may 
carry a quarter of a mile" (N. B. Green, W. Va., 1938, pp. 79-80). 

Breeding: March-July. Amplexation axillary. Eggs in masses, 10-50, at- 
tached to vegetation or trash. Egg complement 300-1500; vitellus 1.6 mm.; en- 
velope 6.0-8.5 mm. Tadpole period 50-60 days. Tadpole 25 mm. Labial tooth 
rows %. Transformation size 8.0 mm. (after Green, Walker, Welter, Carr, 
Barbour, and Sollberger, the last four of whom have sent us many a puzzling 
specimen of this species, ot Hyla cructfer, and of Acns). 

"Its breeding season begins somewhat later than that of trisenata and is more 
prolonged. Clasping pairs and fresh eggs have been found as early as March 
2oth and as late as May i2th. In the same region our dates for tnsenata eggs 
range from March i4th to April i6th. The eggs of brachyphona are so much 
like those of tnsenata and arc laid in similar masses, attached to twigs, leaves, 
or grasses, often well below the surface of the water" (C. F. Walker, Ohio, 

932, P- 3 8 3)- 

"Egg masses were found in the tegion under observation (Randolph and 
Tucker counties, West Virginia, at an elevation of 2000 feet) on April 5. 
Walker (1932) states that iresh eggs have been found as early as March 20, 
presumably in southern Ohio. Pairs in amplcxus and eggs were taken at Por- 
terwood, Tucker County, on June 10, 1936, at an elevation of 2200 feet and on 
Point Mt., Randolph County, on June 7, 1937, at an elevation of 3500 feet fol- 
lowing a rainy spell of several days. Freshly laid eggs were collected by C. J. 
Coin at Camp Woodbine, Nicholas County, on July 2, 1936, at an elevation of 
2000 feet. His notes for this date read, 'Small pool visited at dusk following a 
rain. A long dry spell had preceded this rain. One clutch of freshly deposited 
eggs was observed. . . .' On April 13, 1937, five females were collected on their 
way to the pools. In the laboratory each female was put in a separate jar with 
a male in an effort to determine the number of eggs an individual laid. They 
laid respectively 1479, 383, 318, and 406 eggs" (N. B. Green, W. Va., 19^8, 
p. 80) . 

"Egg masses were collected from a conical hole about two feet deep in the 
vicinity of Reightown on April 22, 1939. On the evening of June 30, 1939 a 
single specimen was collected in a ditch in a field adjacent to the highway be- 
tween Homer and Ruggles Gap" (H. D. Yoder, Pa., 1940, p. 92). 

"A very common spring frog. Breeding begins as early as February 12 when 
the marshes ring with the songs of this species and of Hyla cructfer. It is as 
abundant as H. critcifer and is usually found with it" (W. A. Welter and K. 
Carr,Ky., 1939,?. 129). 



231 HANDBOOK OF FROGS AND TOADS 

Journal notes: July 18, 1931, Beckley, W. Va., Fair Grounds woods. The 
botanists found a queer Psettdacns jenantm near a sawdust pile. 

July 19, Beckley, W. Va., Fair Grounds. Tonight after a heavy ram, in a 
puddle beside the grandstand and in pools beside the track heard several Hyla 
versicolor, and in the distance several Hyla crucijer, but the notes which drew 
us to the spot were new surely a Pseudacris in character of voice, not a Hyla 
crucijer nor an Acris. It was different from Pseudacris n. tnscriata and Pseuda- 
cns n. jenantm. When we first caught it, its long legs reminded us of an Acris, 
and we looked for the rear femoral stripes. They were not present. It seems like 
a Pseudacris, but has too large disks for a normal Pseudacris species. Must look 
up Cope's P. j. brachyphona. 

July 21. Prof. P. C. Bibbee, of State College at Athens, Mercer Co., brought 
us a jar of preserved swamp cricket frogs. They have such queer coloration 
(more like an Acns type) and such spindling legs. Are they a Hyla, Pseudacris, 
or Acris? They must be Pseudacris. Bibbee let me take a few for our collection 
and said they collect them in the spring. Surely someone will have to collect 
Pseudacris across southern Pennsylvania from Carlisle (type locality of P. jeria- 
rum) to southwestern Pennsylvania and West Virginia to see where this form, 
if P. f. brachyphona, meets P. fenarum jenantm 9 m mid-Maryland or mid- 
Pennsylvania. 

July 24. Visited R. K. Brown's laboratory at Blucficld, W. Va. He had a jar 
with several puzzling Pseudacris. 

C. F. Walker was the first author to redescribe Cope's Chorophilus jenarum 
brachyphonus t and it is interesting that four groups at about the same time 
independently concluded to revive Cope's description; Walker on Ohio ma- 
terial, Netting on west Pennsylvania and northern West Virginia material, 
Dunn on west Maryland and Pennsylvania specimens, and we on southern 
West Virginia material. But the question is not entirely settled yet. Is this form 
in eastern Tennessee ? Even as early as 1931 or 1933 when this form was re- 
vived, we had seen material from northern Mississippi and northern Alabama 
to cause us to credit it to those states. Viosca (Ala., 1938, p. 201) furnished a 
very fortunate record. 

A former student supplies the following: "The surprise of the trip was a 
voice heard for the first time near Moundville. The origin of the strange 
'Qurack,' suggestive of the voice of Hyla squirella, eluded us for about twc 
hours before the first specimen was located; then the next six, all males 
were taken in about 15 minutes. This locality, in the foothills at the extreme 
southwestern limits of the Appalachian Highlands, extends the range of this 
species \P. brachyphona} some 500 miles southwestward" (letter). "To deter 
mine the number of eggs laid by each female, ten average-sized individuals 
were dissected. The largest number of eggs found in any one of these was 1202 
and the smallest number was 983. The average number found in ten female* 
was 1092 eggs. The eggs were deposited in masses attached to a stem or leaf. 



TREE FROGS: HYLIDAE 




Plate XLV1I. Pxudacris brachyphona Plate XLVIII. Pseudacris brimlcyt (X 

XO. 1-7- Males. i,2,4,5> 6 - Males - 3,7- Females. 



234 HANDBOOK OF FROGS AND TOADS 

The number of eggs found in each of eleven masses was counted, and the num- 
ber found to vary from 28 to 40, the average being 34 per mass. In the labora- 
tory at temperatures varying from 1 8 to 22 degrees [C|, hatching occurred 
from 72 to 96 hours after the eggs were laid. In the pools, however, where the 
temperature ranged from 2 to 13 degrees, hatching did not occur for a week or 
ten daysS. Several tadpoles kept in a large aquarium in the laboratory at room 
temperature metamorphosed from forty to fifty days after hatching. Tadpoles 
in outdoor pools metamorphosed in forty-five to fifty-five days" (R. W. Bar- 
bour and E. P. Walters, Ky., 1941, p. 116). 

Authorities 9 corner: 
C. F. Walker, Ohio, 1932, pp. 380-381. 
P. Viosca, Jr., Ala., 1938, p. 201. 
N. B. Green, W. Va., 1938, p. 79. 

Brimley's Chorus Frog 

Pseudacns brimleyi Brandt and Walker. Plate XLVIII; Map 16. 

Range: Dismal Swamp, Va., south in the Coastal Plain to Bryan Co., Ga. In 
North Carolina, Brandt has taken bnmleyi in Beaufort, Greene, Pitt, and 
Wilson counties, and has voice records from Craven and Edgecomb counties. 
Newbern, N.C. Type locality Washington, N.C. (after Brandt and Walker, 
N.C., 1933, p. 5). Charleston, S.C. (E. B. Chamberlain, S.C., 1939, p. 17). 

Habitat: Heavily wooded flood-plain pools. Swampy areas. Brandt (N.C., 
J 93^b, p. 500) called it a "palustrine species. . . . This chorus frog was selected 
as the representative of an arboreal form which has reverted to terrestrial life." 
"It is of especial interest in comparison with its more arboreal relative of simi- 
lar size, Hyla crncifer. About four months are spent at the breeding pools; the 
remainder of the year this species usually spends along swampy shores." 

Size: Adults, i-iVi inches. Males, 24-28 mm. Females, 27.0-32.5 mm. 

General appearance: "A dark middorsal stripe from tip of snout to vent. On 
each side of this a similar stripe extending from eyelid to posterior end of 
trunk. A darker, sharply defined line from snout through eye, and along side 
to groin. Dorsal surfaces of legs with several large dark spots, the long axes of 
which are longitudinal, and many smaller dark spots. A distinct dark line 
along outer edge of tibia and on under side of foot. A light line along upper 
jaw continuing back to shoulder. Edge of jaw dark. An irregular, more or less 
discontinuous, dark line along posterior side of arm. Under surface light in 
color with dark spots on chest. Smaller but similar spots on under surface of 
legs" (B. B. Brandt and C. F. Walker, N.C., 1033, P- 3) 

"A medium-sized Pscndaeris with long legs and a trheriate dorsal pattern, 
closely related to the mgritajerianim-tnseriata-brachyphona series, but differ- 
ing in having a more delicate, smoother, skin; a sharply defined black lateral 
stripe combined with a rather weak dorsal pattern; a pronounced tendency 



TREE FROGS: HYLIDAE 235 

towards longitudinal rather than transverse leg markings; a narrow dark line 
along outer edge of tibia; well defined dark spots on chest in most individuals; 
and in life, a distinctly yellowish venter" (same, pp. 2-3). 

Color: "Dorsal color brown (yellowish to reddish shade) ; dark mid-dorsal 
stripe from snout to vent, and on each side of this a similar stripe from eyelid 
to posterior end of trunk. A darker, sharply defined lateral line from snout 
through eye to groin. (In light phases the dorsal stripes are usually obscure, but 
the lateral stripes remain black and sharply defined.) Under surface yellowish 
with small dark spots" (E. B. Chamberlain, S.C., 1939, p. 17). 

"Living specimens show considerable variation in the ground color of the 
upper parts, but are apparently restricted to yellowish and reddish shades of 
brown, ranging from a pale buffy brown to a very dark shade near Light Seal 
Brown (Ridgeway). The dorsal stripes and the peculiarly shaped leg spots are 
usually obscure in the lighter phases, the lateral stripes remain black and 
sharply defined. The color of the venter also varies but seems always to be 
yellower than that of mgrtta. The iridescent surface makes an accurate descrip- 
tion of this color difficult. In the paler individuals it approaches Cream-buff, 
and in more richly colored examples it is near Buff-yellow or Pale Orange- 
yellow. The light cheek stripe below the vitta is similarly colored. Frequently 
there is a deeper yellow spot in the groin. The undersurface of the legs is gray- 
ish. The scattered dusky pigment of the gular sac is underlain with greenish 
yellow. There is an obscure dark bar through the eye. The iris shows gold and 
reddish flecks, the latter restricted to the lower half" (B. B. Brandt and C. F. 
Walker, N.C. and Ga., 1933, p. 4). 

"The more distinctive marks arc the black lateral stripe which extends 
through the eye almost to the groin, the longitudinal not transverse markings 
on the hind legs, and the spotted under parts" (C. S. Brimlcy, N.C., 1940, p. 18) . 

Structure: Smoother than P. n. ntgrita. Venter coarsely granulated; heel to 
anterior eye; disks slightly dilated; muzzle rounded and projecting; tympa- 
num distinct, smaller than eye. 

Brimley (p. 15) gives these characters: "Upper parts with small warts at least 
on the side; snout longer than width across lower jaw at base; snout rounded 
when viewed from above; breast spotted." 

"The leg markings of brimleyt and oculans remain quite different. An ap- 
proach to the longitudinal spots of bnmleyi is seen occasionally in brachy- 
phona, and the voice also resembles that of brachyphona, which is a very dif- 
ferent frog in head shape, pattern, and skin texture" (B. B. Brandt and C. L. 
Walker, N.C. and Ga., 1933, p. 6) . 

Voice: "Walker, who heard the species in northern Georgia, considers the 
call very similar to that of Pscudacris brachyphona. To the writer the call 
sounds like a rasping trill, 'Kr-r-r-ak/ somewhat less than a second in duration. 
The note is suggestive of that of Hyla squirella but is more strongly accented 
at the end, and the intervals between the individual calls are shorter. Calling 



236 HANDBOOK OF FROGS AND TOADS 

has been heard in rare instances as early as November" (B. B. Brandt, N.C., 
19363, p. 218). 

"The type locality is Washington, N.C., where it is the earliest breeding 
species of the genus, mated pairs having been taken by Brandt and his friends 
of the Washington Field Museum from February 19 to April 17, 1933" (C. S. 
Brimley, N.C., 1940, p. 18). 

We still need detailed descriptions of the egg masses, individual eggs, tad- 
poles, and transformation stages. 

Journal notes: Feb. 22, 1934. We have visited the Bughouse at Washington, 
N.C., and George Ross has shown us some of the live adults and tadpoles of 
P. brimleyt. There are yet no detailed descriptions of the eggs and tadpoles, 
but they do not depart markedly from the Pseitdacns type. 

Feb. 2$, 1934. Left Kmston, N.C., southward on No. 1 1 at 2:30 P.M. Farther 
along in a cattail and roadside ditch heard a few isolated Pscudacns. Are they 
Brandt's P. brimlcyi or what ? 

Authorities 9 corner: 
B. B. Brandt and C. F. Walker, N.C. B. B. Brandt, N.C., 1936^ p. 525. 

and Ga., 1933, p. 5. E. B. Chamberlain, S.C., 1939, p. 17. 

B. B. Brandt, N.C., 19363, p. 218. 

Swamp Cricket Frog, Swamp Chorus Frog, Swamp Tree Frog, Swamp 

Tree Toad, Striped Tree Frog, Rough Chorus Frog, Le Conte's 

Chorus Frog, Black Chorus Frog 

Pscudacns nigrita nigrita (Le Conte). Plate XLIX; Map 17. 

Range: North Carolina to northern Florida. Mississippi, possibly into Loui- 
siana. The western limits are not very clearly defined. 

Habitat: Breeds in ditches, ponds, or bayous, and later moves to dryer ham- 
mocks and ridges of the pine barrens. 

Size: Adults, %-iH inches. Males, 21-28 mm. Females, 22-30 mm. 

General appearance: This small frog has a slender body and pointed snout, 
prominent eyes, and long legs. The skin is finely granulated. This form has the 
most numerous small dorsal spots of any Psettdacns. It is gray or olive to black 
in color, with three irregular rows of dark spots on the back and darkly mot- 
tled sides. The dark spots are outlined with white dots. The row of dark spots 
down mid-back divides into two rows toward the rear. A light line is present 
along the jaw with a dark area extending from the snout through the eye and 
beyond the tympanum. There is a light speck back of the tympanum. The legs 
are barred with black spots. There is no triangle or crossbar between the eyes. 
The male has a dark greenish yellow throat with longitudinal folds in the 
middle. 

Color: Female. Okefinokee Swamp, Ga., June 15, 1922. Back dark gray, 
smoke gray, drab, or grayish olive. From snout down middle of back is one 



TREE FROGS: HYLIDAE 



237 




238 HANDBOOK OF FROGS AND TOADS 

row of more or less connected dusky drab or deep brownish drab to black 
spots. Stripe along jaw pale rose purple, pale salmon color, or cartridge buff 
with some seafoam green. 

The row of spots continuous down the back becomes at halfway point two 
rows of separate spots. On each side is one more row of separate spots. Dark 
area from snout through eye, back of eye, and extending along the side to 
groin. Crossbars of legs are same color as spots of back, and light thin bars be- 
tween the spots of legs are like the background of the back. Under parts white. 
Iris black with brownish vinaceous dots. At times, iris has considerable glass 
green. (See A. H. Wright, Gen., 1932, pp. 199-200 for various color character- 
izations.) Sometimes the frogs are blackish olive, and white or creamy below. 
The maxillary stripe may be white, greenish, or a little yellow. 

Structure: Slight webs at bases of second, third, and fourth toes; slight disks 
at tips of fingers and toes; hind legs long; in many ways this is the most dis- 
tinctive of all six subspecies of P. mgnta. 

The original description "Above black speckled with small white warts; 
middle of back cinereous with an interrupted stripe of black, upper lips with 
a white line; beneath granulate whitish; irides golden; legs barred with whit- 
ish, hind part of thighs brown; hind legs very long" (J. Le Conte, Gen., 1825, 

P . 282). 

Voice: "Its call is similar to that of the Cricket 'frog,' but much louder, and 
the crepitations arc slower" (R. F. Deckcrt, Fla., 1915, p. 23). 

In 1921 at Fargo, Ga., F. Harper noted in his journal: "During downpours 
on the afternoon of August 5 ... began to hear several frogs calling in a field 
grown with grasses and sedges and flooded with rain. The notes consisted of a 
shrill metallic staccato trill, frequently repeated: i-i-i-ik, i-i-i-ik, i-i-i-ik. They 
were indistinguishable from those of representatives of this genus in the Atha- 
baska region (septentnonahs) and in the District of Columbia (feriantm)" 

Breeding: Breed from December to April. Larval period 40-60 days. Trans- 
formation April i-June i or July. Size about 9-15 mm. 

Several naturalists in the South have made notes on the breeding of this early 
species. At Biloxi, M. J. Allen (Miss., 1932, p. 7) wrote: "Several pairs were 
taken in copulation on Dec. 15, 1929 and forty-two breeding pairs and some 
clumps of eggs in daylight on January 20, 1931." 

Carr recorded: "Breeding: Jan. 16 to April 2, in temporary flatwoods ponds 
and ditches and in the shallow margins of cypress ponds. January and Febru- 
ary choruses almost always in company with those of P. ornata. The young 
emerge in April and May. They often remain for several weeks about the 
ponds from which they emerged" (A. F. Carr, Jr., Fla., 1940, p. 56). 

Journal notes: June 15, 1922, Folkston, Ga. Near Chesser Schoolhouse under 
chips, small boards, and logs found three Psetidacns in dry pine barrens, as- 
sociated with Bufo qtiercicus. Area beneath boards and cover more or less dry. 



TREE FROGS: HYLIDAE 




Plate XLJX. PseudacrU nigrita nigrita 
XO. i-4 Adults. 



Plate L. Pseudacns ntgnta dar\ii (XO- 
1,3,7. Females. 2,6. Males. 4. Eggs attached 
to small twig. 5. Tadpole. 



2 4 6 HANDBOOK OF FROGS AND TOADS 

July 15, Folkston, Ga. On high sandy ridge south of Spanish Creek, Quercus 
catesbaei, Quercus cinerea, Gaylussacia dumosa and a very much serrated poi- 
son ivy (Rhtts) were growing. Miles and I were looking at the Rhus carefully 
and gingerly. He saw something jump among it. Thought it a grasshopper but 
he soon found it to be a frog. It was Pseudacris n. nigrita. We took its picture 
right there. Other plants there were persimmon, Baptisia, brake, and croton. 
The frog was first on a dead twig of a fallen branch. It would leap among dead 
leaves, among wire grasses, and among the ivy. 

Aug. 16. In making a detour on the Dixie Highway about a mile south of 
Hilliard, Fla., we came to the road opposite a wet pine woods. In one little 
temporary pond in a tussock of grass beside the small pool was a Pseudacris 
n. nigrita calling. To me this form which was finally captured sounded like 
our New York Pseudacris. To F. Harper it sounded like a metallic staccato 
trill, five notes, ic, ic, ic, ic, ic. All our records in July and August are clearly 
sporadic croakings long after breeding and only after heavy rainfall. 

Authorities 9 corner: 

R. F. Deckert, Fla., 1915, p. 23. E. B. Chamberlain, S.C., 1939, p. 15. 
C. S. Brimley, N.C., 1940, p. 18. 

Clarke's Chorus Frog, Clarke's Striped Tree Frog, Striped Tree Frog, 

Striped Chorus Frog 

Pseudacris nigrita clarion (Baird). Plate L; Map 18. 

Range: Texas to Kansas. "The extension of the range of the species north- 
ward has been a gradual and a continuous one, with the proper interpretation 
of the recorded species, so that the discovery of the species in Kansas is not so 
surprising. Cope extended its range to northern Texas (Dallas) in 1880, and 
into Oklahoma in 1894, shortly after recording it from the panhandle of Texas 
(1893). Ortenburger then (1926) reported nigrita (almost certainly clar\ii, by 
this interpretation) from Creek county, Oklahoma, a record verified by a speci- 
men from the same locality in the K. U. Museum, collected by Dr. Taylor. 
Burt (1932) found it in Kay county, Oklahoma, and subsequent collecting 
and examination of the K. U. material has revealed its existence far northward 
into central Kansas, in essentially the same ecological relations as exist where 
it is found in central Oklahoma" (H. M. Smith, Kans., 1934, p. 465). 

Habitat: Abundant in vicinity of marshes. Breeds in roadside ditches, shal- 
low water-lily ponds, shallow mesquite ponds, grassy ponds, or other transient 
pools. 

"Chorophilus tn*eriatits Wied. . . . Baird's types of Helocoetes clarion were 
from Indianola. In East-Central Texas, during the breeding season, this species 
fairly swarms in the roadside ditches and in shallow pools on the grassy flats" 
(J. K. Strecker, Jr., Tex., 1915, p. 48). 



TREE FROGS: HYL1DAE 241 

"Chorophdtts trisenatus clarion. Baird and Girard. ... An abundant spe- 
cies, especially so in the vicinity of marshes" (J. K. Strecker, Jr., Tex., 1902, 

P-7)- 

"Abundant and noisy in pools near the Cimmaron River, at Tucker, Okla- 
homa. Similar to the individuals reported by me from Clarendon, Texas" 
(E. D. Cope, Okla., 1894, p. 386). 

"The specimens found in Oklahoma were in wooded areas. They seemed to 
avoid the deeper, muddier, less protected pasture pools where Scaphiopus and 
Bufo were numerous. Some were found, however, hopping about m the grassy 
fields, not far distant from pools. The specimens from Sedgwick county, Kan- 
sas, were found in a habitat essentially similar to the above, except that the pool 
in which they were found was permanent, while the other apparently was not" 
(H. M. Smith, Kans., 1934, p. 464), 

Size: Adults, %-i!4 inches. Males, 20-29 mm - Females, 25-51 mm. 

General appearance: This is a small grayish or olive frog with dark longi- 
tudinal spots, or these spots may be arranged m three stripes. A dark lateral 
banding extends to the nostril. There is a light stripe on the upper jaw. The 
legs are barred above and white or pale buff beneath. Venters arc white or 
ivory yellow with the throat of the male a dark olive buff. They are protectively 
colored little things in the grassy edges of the ditches where they call and breed. 

Color: Four males. Brownsville, Tex., Feb., ig2$. Upper parts light grayish 
olive, pale olive-gray, grayish olive, or tea green. Spots on back and triangular 
spot and bars on legs yellowish oil green, oil green, or courge green. Bar from 
snout through nostril, through eye and tympanum, and onto sides black. 
Sometimes back of shoulder it has the greenish color of dorsal spots. Triangu- 
lar area between eyes equilateral in one, long-caudally prolonged in another, 
divided into three spots, the two forward angles each making a spot and the 
caudal angle prolonged backward. Area in another not a triangle but with pro- 
longation backward at one end like a figure seven. Two of these four have a 
spot ahead of the triangle. Above lateral vitta, through eye, over shoulder is a 
parallel series of dorsal spots, usually two, the caudal one longer, or the cephalic 
one larger, or all one or occasionally more than two. If there are other dorsal 
spots, they are in the posterior half of the back. There is no distinct middorsal 
line of spots or line. Belly and under parts white except for throat, which is 
olive-ocher, orange-citrine, olive-yellow, or lemon chrome. 

Detailed color descriptions of a pair taken at Bceville, Tex., March 26, 1925, 
follow: 

Female. Back deep olive-gray; on sides pale olive-gray above lateral stripe. 
Spots on back hellebore green surrounded by black. Stripe through eye and 
along side dark grayish olive. Bars and spots on fore limb like lateral stripe. 
Stripe on upper jaw cartridge buff with marguerite yellow or primrose yellow 
in it. Tympanum brownish olive or light brownish olive. Venter white, pale 



2 4 2 



HANDBOOK OF FROGS AND TOADS 



olive-buff, or ivory yellow. Underside of hind legs light grayish vinaceous or 
light brownish drab. Iris upper part of rim marguerite yellow or primrose 
yellow, rest dotted black, primrose yellow, and cinnamon drab. 

Male. Backgiound pale smoke gray, also areas of tea green. Spots vetiver 
green, surrounded by black. Lateral stripe bone brown. Tympanum army 
brown. White or primrose yellow spots on tubercles below vent and on basal 
insertion of thighs. Throat discolored dark olive-buff or more yellowish. Two 
longitudinal plaits in middle of throat. Iris of this male has also some avella- 
neous in it and a tendency to a dark horizontal bar through it to complete nasal 
and lateral stripe. 

March 25, 1925. In the two pairs which laid, the females were more spotted, 
the males more striped. Is this a natural tendency or an individual variation? 
The male of a third pair is veiy much striped. Another male we captured is 
very much spotted. 

Structure: Snout acute, projecting beyond lower jaw; toes slightly dilated at 
tip^; male throat with one to three longitudinal folds. The hind limb, tibia, 
snout, and head are longer than in P. n. tnsenata. 

The original description "Helocoetes darken Baird. Snout acute, projecting. 
Extremities somewhat dilated. Tibia half the distance between eye and anus. 
Foot but little longer, not nearly half the length of body. Above grayish brown 
or ash, with distinct large circular blotches. A dark band from snout through 
eye and tympanum down the sides, and a whitish line on the side of jaw. Size 
about one inch long. Hub. Galveston and Indianola, Texas" (S. F. Baird, Gen., 
1854, p. *j). 

Voice: The call is a grinding note, more measured, and not so shrill as the 
call of Acns. It is raucous and grating, it-it I it-it /it-it, sometimes 60 times with- 
out a stop, sometimes uniform, sometimes double speed. They call from under 
the grass edges. Sometimes there is a synchronized chorus of sawing notes. 

March 24, 1925, Beeville, Tex. Air is resounding with Pseitdacrts. 8:30 P.M., 
air 74. Croaked 24 times, stopped instant, then began again 25 to 30 times, 
hardly catch a breath then start again, another 29 in 30 seconds. They are hid- 
den under the grass edges like Mtcrohyla, which is calling at the same time in 
the same roadside ditches. It is the dominating note over Mtcrohyla and Bufo 
valliceps just now. 

"Sometimes in later afternoon or early dusk, preceding a heavy chorus dur- 
ing the night, P. clarion males call as they approach a breeding pool after the 
manner of Hyla v. vemcohr. These calls, however, are enough different from 
the typical breeding cry that when I first heard them I was not sure of the spe- 
cies responsible till I had run down and captured a specimen. Such calls are 
not so loud nor so frequent as the typical breeding call. They have an indescrib- 
able quality which for want of a better term, I call 'pensive' or 'pleading.' Be- 
cause of these differences I suspect (but have not proved) that this pensive call 



TREE FROGS: HYIJDAE 243 

functions to attract other males to appropriate breeding pools that have been 
sensed by males nearer to them. At least, such calls in my experience have been 
limited to a circle of some four hundred feet in diameter whose center is cov- 
ered by water appropriate for breeding activities. How the first males find a 
pool in this (and other) species is almost entirely unknown" (A. N. Bragg, 
Okla., 1943^ pp. 131-132). 

Breeding: The species breeds with the spring rams, March 5~May 20, occa- 
sionally in summer to mid-August. The citrine drab and ivory yellow eggs are 
in a loose irregular mass attached to plant stems, their number few, 150 to 175 
eggs in groups of 3 to 50. The egg is %2-/r> inch (0.65-0.9 mm.), the outer 
envelope M 2-M n inch (2.2-2.4 mm -) or more loose and irregular, the inner 
envelope K inch (1.4-1.8 mm.). The grayish olive tadpole is small, TH-I/^ 
inch (25-30 mm.), its crests nearly transparent, the tooth ridges %. After 30 to 
45 days, the tadpoles transform, April i to June 20, at r 'i<j-/{> inch (cS-i 3 mm.). 
H. M. Smith's (Kans., 1934, P- 4^4) specimens were slightly larger, 13.5-17.0 
mm. in length. 

One pair in captivity laid 154 eggs in several small masses arranged about 
one inch to one and a half inches apart on the stem. These masses were 17, 27, 
30, 37, 16, 13, and 14 eggs, respectively. The last mass rested on the debris in 
the bottom of the jar. There is a very loose outer envelope, at times irregular. 

We found the tadpoles transforming in extreme southern Texas on April 22. 
From the region of San Antonio we have the following note: May 29, 1925. 
Some Psetidacris are approaching the fully developed two-legged stage. Si'/c, 
8-13 mm. 

"Found breeding in temporary pools from early May to mid-June. Com- 
mon" (G. A. Moore and C. C. Rigney, Okla., 1942 [1941 1, p. 78). 

"Feb. 25, 1944. Heard Pseiulacns mgnta clarlyi frogs at Duck Pond about 
2^/2 miles north of my station. These arc first near here. Formerly nearest were 
near Lytle (9 miles)" (A. J. Kirn, Tex.). 

Journal notes: March 25, 1925, Beeville, Tex. The pair we caught last night 
broke and one was lost. Captured an extra female. Put in photographic jar, two 
females and one male. This morning when I arose, they were mated, but had 
not laid. The female of this pair was not the female of the original pair cap- 
tured. They laid in one-half hour or less. The first pair captured had been rest- 
ing at the grassy edge of a roadside ditch. The female was larger than the male. 
The embrace was axillary. 

April 7, 1925. The eggs laid in Bccville, March 25, are developing nicely. 

April 22, San Ben i to, Tex. In sedges around a pond one mile south of San 
Benito, these little transforming frogs were swimming in shallow water. They 
often duck. Hard to collect without killing them, they are such delicate little 
creatures. Skinned hind legs of two in their capture. They can turn heads like 
some other Psettdacris. Found only a very few tadpoles in the pond. The species 



244 HANDBOOK OF FROGS AND TOADS 

must be through metamorphosis in this pond. This is a beautiful water-lily 
pond with Marsilca and Castaha elegans. 

Authorities' corner: 
E. D. Cope, Tex., 1893, P- 333- 
J. K. Strecker, Jr., Tex., ipioc, p. 80. 
A. N. Bragg, Okla., 1941, p. 52. 

"The comparison of these two tadpoles \triseriata and darfyi] reveals an in- 
teresting situation. They arc so nearly alike that those who believe them of 
separate species (e.g., Smith 1934; Bragg 1943) can take little comfort from the 
data. However, those who call them subspecies or varieties of one species (Burt, 
1932; Wright and Wright, 1935, 1943; Stejneger and Barbour, 1939) must ex ' 
plain the proportional differences reported. It would seem, therefore, that either 

(1) one of these forms has fairly recently been derived from the other, or 

(2) both have been derived in recent geological times from a single ancestral 
form. The latter seems the more probable in view of lack of intergradmg speci- 
mens where their ranges meet" (A. N. Bragg, Okla., I943C, p. 138). 

Having seen most of the tadpoles of the United States, even though we have 
not described them all minutely, we smile at "take little comfort from the 
data." We would have faith today in no one who pretended he could distin- 
guish the tadpoles of all Psettdacris or all Bttfo species. Bragg, than whom no 
one has been more active recently in life histories, must have enjoyed writing 
the paragraph above. P. n. clarfyi and P. n. triseriata are close. When we find 
several times in the field (not in alcohol) pairs of P. n. clarfyi with the males 
striped and the females spotted, or when we find males longer-legged and fe- 
males shorter-legged in some P. n. septentnonahs specimens, it means that 
herpetologists do not know the species; or original descriptions alone do not 
solve; or that the Pseudacris nignta complex is difficult to explain by any classi- 
ficatory schema conceived long ago. 

Eastern Chorus Frog, Eastern Swamp Cricket Frog, Swamp Tree Toad, 
Swamp Tree Frog, Chorus Frog, Striped Tree Frog, Common 

Chorus Frog 

Pseudacris nignta jenartim (Baird). Plate LI; Map 17. 

Range: New Jersey, Pennsylvania to North Carolina, and possibly into 
South Carolina. 

Habitat: Breed in marshy stretches or shallow ponds, or at edges of ponds 
where there is matted vegetation, sedgy clumps, or moss. 

"Between the railroad station at Annadale and Seguine's pond [Staten Is- 
land j, there are several marshy places and one little artificial pond. For several 
years we have heard in these lowlands, from the middle of March to the first 
week in May, the song of what we considered to be the swamp tree frog, Cho- 
rophilus triseriattts (Wied)" (W. T. Davis, N.Y., 1912, p. 66). 



TREE FROGS: HYLIDAE 245 

"This species I have found abundant on the sides of pools and ponds in the 
neighborhood of Gloucester, N.J., in the spring and early part of summer. It 
delights in those small and often temporary pieces of water which are inclosed 
in the densest thickets of spiny Smilax and Rttbtts, with scrub oaks, and sur- 
rounded by the water loving Cephalanthus, where no shade interrupts the full 
glow of sunlight" (E. D. Cope, Gen., 1889, p. 344). 

"As the pools in which they stay are usually full of grass, low rushes and 
other vegetation, they are not easy to catch although patience and perseverance 
will secure specimens without too much trouble. . . . Outside the breeding 
season these frogs are not often seen, but may be sometimes found hopping 
about in damp places or in cool, shady woods. On the ground they are not 
much more active than toads" (C. S. Bnmley, N.C., 1940, p. 18). 

Size: Adults, %-i% inches. Males, 21-30 mm. Females, 22-^ mm. 

General appearance: These are small frogs with pointed heads, extremely 
long fourth toes, very little or no webbing between the toes, and with small 
disks on their toes. They are grayish olive or brown. There is a dark triangle 
between the eyes that extends backward as a dorsal stripe, with another stripe 
on either side. A dark stripe extends through the nostril and eye to the shoul- 
der. This is bordered below with a light stripe on the upper jaw. The throats 
of the males are usually dark but not always so. The under parts are a buffy 
cream color. 

Color: Carlisle, Pa. Male. Back deep grayish olive, drab, or drab-gray. Tri- 
angle between eyes going back as dorsal stripe with a stripe on either side. Each 
of these may be light-centered with body color. These dorsal stripes may be 
black, grayish olive, light grayish olive, or deep olive. Undersides of hind legs 
are brownish drab or benzo brown. Belly cartridge buff, cream-buff, or cream 
color. Throat buckthorn brown near tip to yellow ocher in posterior part. Band 
from snout to halfway along sides mummy brown or black. Ins same, with 
little burnt sienna in upper half. Stripe along upper jaw almost to shoulder, 
warm buff, cream-buff, or chamois. 

Female. Upper part of head, back, forelegs, and hind legs, russet, hazel, 
verona brown, or warm sepia. Underside of hind legs benzo brown, drab-gray, 
or brownish drab. Under parts of body cream color, becoming slightly massicot 
yellow or naphthalene yellow on throat. Stripe down middle of back, stripe 
either side of median stripe, and faint spots on sides arc black, chestnut brown, 
mummy brown, or sepia. Stripe through nostril and eye to shoulder and be- 
yond warm sepia or Vandyke brown. Band around upper jaw extending back- 
ward beyond shoulder below the dark stripe is warm buff, cream buff, or 
chamois. Iris same color as vitta through eye but dotted with mars orange or 
burnt sienna. These predominate in the upper half, giving the iris a "reddish" 
or "coppery red" appearance. 

Structure: The original description "Helocoetes feriarum, Baird. Body 
stout, squat. Head broad. Femur and tibia and hind foot about equal, and half 



246 HANDBOOK OF FROGS AND TOADS 

the length of the body. Above dark or fawn, with three nearly parallel stripes 
down the back, the central widening, but scarcely bifurcate behind, and com- 
mencing behind a triangular spot between the eyes. A similar dark vitta on 
sides of head and body, with a white line along edge of the jaw. Body about one 
inch long. Hab. Carlisle, Penna." (S. F. Baird, Gen., 1854, p. 60). 

Voice: A penetrating and grating it-it/ it-it is given many times without a 
stop. "Its note resembles that of Acns in being crcpitant, and differs from the 
toned cry or whistle of the Hylae. It is not so loud as the former and is deeper 
pitched; it may be imitated by drawing a point strongly across a coarse comb, 
commencing at the bottom of a jar and bringing it rapidly to the mouth; or 
better, by restraining the voice to the separate vibrations of the vocal cords, and 
uttering a bar of a dozen or twenty vibrations, beginning with the mouth closed 
and ending with it well opened" (E. D. Cope, Gen., 1889, p. 345). 

"This species is rarely seen except during its breeding season, when it is quite 
common. Its cry consists of two or three clear whistlchke chirps, like those of 
a young turkey, and after one has learned to distinguish it from that of H. pid^- 
eringi, serves to betray its presence at once. In the spring of 1896 I collected 
thirty specimens in less than an hour from the gutters along the Conduit Road 
near Cabin John's Bridge" (W. P. Hay, D.C., 1902, p. 129). 

"The earliest of our frogs to breed at Raleigh, and its lazy chorus may be 
heard any warm day in February and March and even in December and Jan- 
uary, though the very early songsters are usually only single individuals. The 
cry is a sort of grating rattle rising at the end, and is uttered by the male frogs 
lying at the surface of some shallow pool. They are very shy and sensitive and 
one always stops singing before an observer gets near enough to perceive the 
author of the noise, while others further away keep up the chorus'* (C. S. Brim- 
ley, N.C., 1940, p. T 8). 

April 15, 1921, Raleigh, N.C. Temperature of air 68F. Tonight at 9, Vernon 
and Julia Haber took me out to the Oakwood Cemetery at East Raleigh and 
to St. Augustine Grounds. Up in the woodland, where we couldn't find it, was 
a Pseudacns fenantm which sounded much like our northern swamp cricket 
frog. In the ponds were tadpoles of the cricket frog in all stages of transforma; 
tion. If anything, these transform at a little larger si/e than those farther north. 
In order of activity the meadow frog and Acns gryllits are practically through 
breeding, though in the ponds the swamp crickets have left for uplands. Fow- 
ler's toads are still in chorus and strongly breeding though started much earlier, 
and Hyla versicolor is just coming to the pond. 

U I have heard the swamps of the barrens and thickets of Southwestern New 
Jersey resound with them as early as the twentieth of March, when a skim of 
ice covered part of the water. I have also heard it in other level parts of the 
same State later in the season, and in the lower part of Chester County, Pa." 
(E.D. Cope, 1889, p. 345). 

Breeding: They breed from February to May 10. The black and cream eggs 



TREE FROGS: HYLIDAE 247 

arc in irregular loose jelly masses attached to the stems of matted vegetation. 
The egg is %s inch (0.9-1.1 mm.), the single envelope %-/ii inch (3.2-4.0 
mm.). The vitelline membrane is close to the vitellus. The envelope in the mass 
becomes larger and irregular. The small tadpole, i-i'Kr* inches (25-33 m m -) 
has the tail medium, with tip acute or acuminate, and is blackish or olive in 
color with bronzy belly. The tooth ridges are %. After 50 to 60 days the tad- 
poles transform until mid-June, at Mc-Vs inch (8-12 mm.). 

April 6, 1929, North Mountain, Carlisle, Pa. There were many egg masses 
attached to the stems in this vegetation mat. The jelly was loose and indeter- 
minate in shape. It stuck to the stems in irregular fashion and to get a mass a 
little chunk of vegetation had to be pulled out. 

"The distribution of Pseudacns is limited to an area in extreme southern 
Berks County from Gerger's Mill almost to Hopcwell [N.J.] and south to 
Elverson. They were found laying eggs at Cold Run, May 4, 1937, by Earl L. 
Poole. Earliest record: calling, Mar. 15, 1939. Full chorus at 45, intermittent, 
scattered calls at 34; an occasional short call at 31; silent at ?oF M (C. E. 
Mohr, Pa., 1939, p. 78) . 

"The eggs arc laid in small masses surrounded by a very weak and watery 
gelatinous envelope and are attached to the stems of plants or other supports 
just below the surface of the water in which they are laid. The tadpoles develop 
very rapidly and I have found them as early as April 16 with the hind legs suf- 
ficiently developed for them to be able to hop although they had not yet lost 
their tails" (C. S. Brimlcy, N.C., 1940, p. 18). 

Journal notes: April 3, 1929. On to Carlisle, through Harnsburg. Just south 
of Harrisburg is a fine grassy shallow pond but no Pseudacns were calling 
there by day. Drove around Carlisle that afternoon to spy out ponds suitable 
for Psetidacris. We heard some out the Mt. Holly Rd. in a punched tussock 
and swampy area in a field. Other ponds between Mt. Holly and Dillsburg 
had many peepers but no Pseudacns. Went out the Chambershurg Rd. (April 
4) and on the way back found a surface pond at the edge of town, in fact in 
a vacant lot near a big building. Heard just three Pseudacns here and caught 
them all. We hated to give up with only three real Carlisle Pseudacns, but the 
next night brought out no more. 

April 6, started up North Mountain. Just before Indian Springs, well up on 
the mountain, we heard a great chorus of Psetidacris feriarum. On the hillside, 
beside the road in an open field was a springy area. Finally we saw two little 
fellows with their amber throats so distended that they showed from the rear. 
They were sitting in the mat of vegetation, a little ways from a tussock, with 
their heads held up vertically, above the water. We went down to catch them 
and found the stream area fairly deep but filled with this mat of grassy vegeta- 
tion. We rolled this back a bit at a time and caught an occasional frog in the 
tangle. One was a female, ripe, and strange to say she looked slender, as did 
the males from the Carlisle Pond, while the males from this Mountain Spring 



248 HANDBOOK OF FROGS AND TOADS 

area were fat and squatty. We caught six or seven males and one female there. 

June 10, 1929. Over the hills from Scranton to Harrisburg, leaving the main 
road at Clark's Ferry, and over the mountain toward Carlisle, back to the spot, 
Indian Springs Garden, where we caught Psetidacris n. feriarum at Easter 
time. Our springy hillside is almost dried up but is still spongy and water stands 
where you step, but no Pseudacris visible. Looked for little transformed ones 
but found none. Bert has gone down to the pond in the meadow below; yellow 
sweet clover, red clover, daisies, wild roses are in bloom. No luck at all. On 
toward Carlisle with crestfallen faces. A pond by the roadside many tadpoles, 
but not the right ones. Came to a small stream with culvert in the road, a deep- 
cut stream with 6-foot banks. Above one bank was a springy inlet with masses 
of sensitive fern, grassy tussocks, touch-me-not, smartweed, and a little sedge. 
To the left in quiet water pockets sometimes no more than 4 to 6 inches deep, 
among grassy tussocks and a little sedge and fern was our big game. Tadpoles 
in general quite dark. As they approach transformation the triangle between 
the eyes becomes quite prominent. It curves in on each side, and the rear point 
extends as a line down the back to the sacral region. On either side of middorsal 
stripe appears a dorsolatcral one of the same length. As the tads rest in the 
water, the tail region seems to be more or less speckled or to have an alterna- 
tion of light and dark. We were impressed with long tail and bronzy belly. 
Some tadpoles were only half grown. 

Authorities 1 corner: 
R. Sutcliff, Gen., 1812, pp. 213-214. 
W. T. Davis, N.Y., 1912, pp. 66-67. 

Queries and comments: i. Good authorities have identified specimens from 
eastern Tennessee, Arkansas, and Illinois as P. n. feriarum, and Del-Mar-Va 
specimens as P. n. triseriata. Are P. n. feriarum and P. n. triseriata one? 

2. In 1921, we became acquainted with P. n. feriarum at Raleigh, N.C., and 
in 1934 we saw in Washington, N.C., live tadpoles and material of P. brimleyi. 
Much remains to be done on these two forms. 

$. Our Carlisle, Pa. (type locality), material did not correspond too exactly 
with Baird's description. 

4. For 30 to 40 years our students from Philadelphia and New York City 
have from time to time queried whether there were two different forms in 
New Jersey. 

5. Is P. n. feriarum coastal in the northern and Piedmont portions and inland 
in the southern portion of its range? 

6. Does P. n. feriarum really reach northwestern Florida? The Tampa Bay 
record must be a mistake. 



TREE FROGS: HYLIDAE 



249 




Plate LI. Pseudacris nigrita feriarum Plate L/7. Pseudacris nigrita septcntrh- 

(XO- 1,2,6- Females. 3,5,7. Males. 4. Eggs. nalis. 1,3,4. Females (Xi). 2. Male (XO- 

5. Female (Xi%). 



250 HANDBOOK OF FROGS AND TOADS 

Northern Striped Tree Frog, Northern Chorophilus, Northern Pseudacris, 
Spotted Tree Frog, Northern Spring Peeper, Peeper Frog, Swamp 

Whistler 

Vscttdacris nigrita septentnonalis (Boulcnger). Plate LII; Map 18. 

Range: Minnesota to Hudson Bay, to upper British Columbia and Macken- 
zie through Montana, Saskatchewan, and Alberta; Canadian Northwest. Ed- 
ward A. Preble found it from Lake Winnipeg to York Factory on Hudson 
Bay and from Edmonton, Alberta, to Ft. Norman west of Great Bear Lake. 
Some authors and collectors have felt inclined to assign some of the swamp 
cricket frogs of our northwest border states from Minnesota to Montana to 
this form, and surely some of the specimens we collected in the Red River of 
the North were short-legged enough to fall withm this subspecies. G. C. Carl 
(B.C., 1945) reports it for Peace River district. 

Habitat: Swampy borders of rivers, lakes, ponds, and meadows. 

Size: Adults, %-i% inches. Males, 19-32 mm. Females, 19-35 ram. 

General appearance: Pembma, N.D. This is a small, long-bodied, short- 
legged tree frog. The snout is pointed. The color may vary from a gray to 
tawny or buff. The most prominent marks are the dark brown lateral stripe 
from snout to groin, and the light creamy stripe on the jaw below the eye. Cer- 
tainly when seen at rest, you are impressed with the very short hind legs in 
sharp contrast with the long body. It has five darker stripes down the body. 
The median dorsal one may be broken into dots, the two laterodorsal stripes 
are more constant and the lateral ones most prominent. The male has a green- 
ish yellow throat, the female, a light throat. The legs of the female arc short 
and stout, those of the male longer and thinner. The under parts are greenish 
white. 

Color: "The following brief description of the color was taken from a live 
specimen from Oxford House: Body light green above, greenish white be- 
neath; body stripes bronzy lavender; tympanum brownish; hind legs light 
green above, flesh color beneath*' (E. A. Preble, Kcew., 1902, p. 134). 

Females. Pembina, N.D., Aug. 30, 1930. First female. The color of top of 
head, top of arms and legs, and back between the three dorsal stripes is drab*. 
The color between outer dorsal stripe and lateral band is a smoke gray or pale 
drab-gray. The band extending from snout through eye and over tympanum 
to near groin is olive-brown or brownish olive. This color is solid to the arm 
insertion, then goes over a pebbly surface, the interstices of which are smoke 
gray or pale drab-gray, like the color above the band. The median and two 
laterodorsal stripes are huffy brown outlined with dotted lines of olive-brown. 
On top of head arc a few specks of olive-brown. The femur and tibia also have 
spots of olive-brown, and on front of tibia and rear of foot and tarsus are half 
bars of the same color. On the upper jaw there is a prominent line of olive-buff 
or chartreuse yellow bordered below, broadly on upper jaw and narrowly on 



TREE FROGS: HYLIDAE 251 

lower jaw, by a finely punctate band of buffy olive or drab. Under surfaces are 
pale dull green-yellow, except for underside of hind legs, which is light grayish 
vinaceous. All five stripes are continuous. The eye is olive-brown, except for 
the upper rim of pupil, which is clear yellow-green; the ins is dotted all over 
with clear yellow-green and mars orange spots; the front, back, and lower mar- 
gin of pupil rim is a succession of clear yellow-green dots. 

Second female. In middle of back the dorsal stripe breaks into two irregular 
rows of small spots. 

Third female. The back is tawny-olive to ochraceous-buff. The dorsal stripes 
are indistinct but nevertheless indicated. The lateral stripe is most prominent. 

Male. Walhalla, N.D., Sept. i, 1950. The area between lateral band and 
laterodorsal stripe is pale olivine or pale fluonte green. The top of back is mi- 
gnonette green. The three dorsal stripes are Kronberg's green surrounded by 
broken lines of olive-brown. The same color extends from snout through eye 
and halfway along the side. The stripe below the eye is glass green. The eye is 
like the lateral stripe in color and has the same dots as the female's, except that 
the upper iris rim is primulme yellow. Under parts are pale niagara green. The 
throat is citrine, old gold, or olive lake. 

Structure: Hind legs short. The original description "Chorophtlus septen- 
tnonalis. . . . Psendacns mgnta, part T, Gunth. Cat. p. 97. Tongue oval, 
slightly nicked. Vomerine teeth in two small groups behind the level of the 
choanae. Head longer than broad; snout subacummate, prominent, twice as 
long as the diameter of the eye; latter very small, can thus rostrahs rather indis- 
tinct; interorbital space a little broader than the upper eyelid; tympanum 
nearly as large as the eye. Fingers and toes moderately slender, latter with a 
slight rudiment of web; first finger shorter than second; an indistinct outer 
mctatarsal tubercle. The hind limb being carried forwards along the body, the 
tibio-tarsal articulation reaches hardly the tympanum. Skin granulate above 
and beneath. Yellowish olive above, with five longitudinal dark bands the 
median bifurcating on the sacral region, the outermost extending from the tip 
of the snout through the eye to the middle of the side; beneath whitish, im- 
maculate. From snout to vent 25 milhm. ... a. 9 Great Bear Lake. Well 
distinguished from C. tnseriatus by its shorter hind limbs" (G. A. Boulcnger, 
Gen., 1882, pp. 335-556). 

Boulenger's key separates this form from mgnta and tnseriata by its rough 
dorsal region. 

Voice: Its call is like those of the more southern forms. 

"This little frog, whose trilling notes arc exactly similar to those of the more 
southern forms of the genus, is fairly common nearly throughout the region 
[Athabaska-Mackenzie Region]. ... At Fort Simpson I first heard its notes 
on May 3, 1904, but failed to take specimens. I heard it also on the Mackenzie 
above Fort Norman early in June" (E. A. Preble, Athab., 1908, p. 502). 

"Though its piercing 'prreep prreep! from the chilly pond, in early spring- 



252 HANDBOOK OF FROGS AND TOADS 

time is familiar to all, very few have seen the originator of the noise or know 
that it is a tiny frog that makes this small steam-whistle. While uttering it, his 
throat is blown out like a transparent bladder, and is nearly as big as himself. 
At Shoal Lake, in 1901, 1 found them still singing in the first week of July. The 
note is more rattled than that of H. crucifer" (E. T. Seton, Man., 1918, p. 83) . 

"Northern Frog. Pseiulacrisigiq: June 18, heard at One Four; Milk River 
July 21, one and again on Aug. 4. Common about McGrath from middle to 
end of August" (M. Y. Williams, Alta. and Sask., 1946, p. 49). 

Harper ( Athab., 1931, pp. 68-70) heard them trilling from May 1 1 to Oct. 21. 

Breeding: This species breeds from May to early June. The eggs arc in an 
irregular jelly mass attached to vegetation. The eggs are Viio-Mo inch (1.2-1.4 
mm.). The tadpoles transform at about %-% inch (7.5-13 mm.). 

In Manitoba E. T. Seton has taken several immature specimens between 
14.5 and 17.0 mm. R. Kennicott at Ft. Resolution, Canada, secured immaturcs 
as small as 16.5-19.0 mm. In Minnesota, in the valley of the Red River of the 
North, the same collector secured immatures 15.0 and 15.5 mm. Young of the 
latter size were secured by J. P. Jensen, September, 1908, at Eagle Bend, Min- 
nesota. In Montana at the Crow Agency M. A. Hanna secured a similar set 
16-18 mm. on July i T, 1916. All these must be well beyond transformation size, 
for three out of the five nigrita subspecies we know transform at 7.5-13 mm. 

Journal notes: Aug. 31, 1930, Red River of the North, Pembma, N.D. Started 
out about 8 A.M. to beat the vegetation along the river bank, expecting to find 
more northern wood frogs. Along the shore is a mud flat, baked and broken 
into squares. The cracks are deep and fairly wide. There are deeper holes 
punched by cows and horses. Many crickets and spiders were running over 
the flats, and clouds of small insects come out of the more moist footprints. We 
walked along the shore, more or less abreast; Bert was in the vegetation and 
I walked on the flats at the edge of the vegetation. I beat the bordering vegeta- 
tion with a cane and searched particularly along the mud at the edge, expecting 
at any moment to sec a wood frog. Suddenly my eye spied a small frog sitting 
at the edge of one of the clumps. As I looked at it, it jumped into a crevice. As 
I looked closely, imagine my surprise to find that it was a Pseudacns n. septen- 
trionalis. We have been looking for suitable places for the search of this specie* 
for some days. Soon after, I saw a second clinging to the edge of another crev- 
ice. As we came close to catch it, it cocked its head to one side as some other 
Pseudacris do at times. A little later, a third appeared clinging to a close-grown 
grassy edge covering clumps bordering the mud flats. All three arc females. 
They certainly have short hind legs. In no case did we get the impression of the 
frog leaping. They must have moved to have caught my eye, but it was not a 
leap like that of a wood frog or meadow frog. The last one acted as if crawling 
up the grass. In captivity they leap from 2 to 8 inches. There are a few meadow 
frogs along the shore and we caught several northern wood frogs. 



TREE FROGS: HYLIDAE 153 

Sept. i, 1930, Walhalla, N.D. The boys in searching the manholes of drain- 
age sewers saw a midget on the side wall of one hole and with searchlight 
climbed down the ladder after it. It proved to be the much-desired game, a 
male Pseudacris n. septentrionalts. It is smaller than the females from Pembina 
and the hind legs look longer and slimmer. 

Authorities 9 corner: 

L. Agassiz, Ont., 1850, p. 379. G. C. Carl, B.C., 1943, p. 46. 

E. T. Seton, Man., 1918, pp. 82-83. W. J. Breckenridge, Minn., 1944, p. 67. 

I. McT. Cowan, B.C., 1939, p. 92. 

Striped Tree Frog, Three-striped Tree Frog, Swamp Cricket Frog, Striped 

Bush Frog, Three-lined Tree Frog, Swamp Tree Frog, Peeper, Spring 

Peeper, Western Striped Frog, Western Marsh Toad 

Pseudacris mgnta tnsenata (Wied). Plate LIII; Maps 17, 18. 

Range: Along the shores of Lake Ontario westward through Ontario, 
Michigan, Wisconsin, Minnesota, North Dakota, Montana, to Idaho south- 
ward to northern Arizona and northern New Mexico. No records in western 
Nebraska and Kansas. Eastern Kansas southeastward through Oklahoma to 
Arkansas and eastern Texas, possibly to northern Louisiana, thence through 
Tennessee and Kentucky (few records) to Ohio and western Pennsylvania. 

Habitat: Low bushes and plants, and on the ground. Breeds in ditches, 
swamps, or temporary ponds. 

"This species, though called a tree-frog, probably never climbs trees. They 
seem to live on the ground among the fallen forest leaves and in the grass" 
(O.P,Hay,Ind.,i8 9 2,p.47i). 

"This is one of the nicest marked of the small frogs, found generally in early 
spring in the ditches along the country roads, Wet Prairie, Madison County, 
111., and near Bluff Lake, St. Clair County, 111. It is often seen during harvesting 
season and for that reason farmers call it the 'Hay Frog' " (J. Hurter, 111., 1893, 
p. 254). 

"This species is almost always found on the ground, where it hides away 
under loose stones or fallen timber" (M. M. Ellis and J. Henderson, Colo., 191 3, 

P. SB). 

"It is to be found in swamps on low herbage or on the ground" (M. Morse, 
Ohio, 1904, p. 119). 

Size: Adults, %-iV> inches. Males, 21-32 mm. Females, 20.0-37.5 mm. 

General appearance: These are small, slender frogs with pointed heads, ex- 
tremely long fourth toes, and small disks on their toes. They are brown, olive, 
or grayish with a dark brown triangle, spot, or stripe between the eyes. The 
most prominent stripe is one from the nostril through the eye, over the arm, 
and extending along the side halfway to the groin. There are three dark stripes 



HANDBOOK OF FROGS AND TOADS 

down the back, continuous or more or less broken up, and dark spots on the 
legs. There is a prominent light cream or silvery line along the upper jaw. They 
are light cream or white beneath. Their skm is finely granular. 

April, 1928, Hilton, N.Y. At night it looks mud-colored. Seldom did the 
stripes or spots appear much revealed. Pointed head, long legs, small and deli- 
cate in form. When perched on a stick or floating mass, sometimes reminds 
me of P. oculans. 

Color: Ann Arbor, Mich., from Mrs. H. T. Gaige, April 7-8, 1929. Female. 
Sayal brown, tawny-olive, or wood brown on back and upper part of fore 
and hind legs and tip of head. Stripes and spots bone brown, black, or chest- 
nut brown. The marks are: a triangle at the eyes extending backward as 
a median stripe, which may be interrupted and resumed and subdivided on 
rear back into two parallel lines; another band along dorsolateral region from 
above shoulder to groin; and a stripe from snout through nostril, eye, and tym- 
panum halfway to groin. Tympanum snuff brown or mikado brown. Stripe 
on upper jaw warm buff or cream-buff with upper part light pinkish cinnamon 
below dark stripe through nostril and eye. Dark edge below this light stripe 
is on labial border. Iris same color as stripe along head, with dottings of mars 
orange or burnt sienna; over pupil rim is light yellow-green. Under parts as in 
P. n. jerarium from Carlisle, Pa., are cream color, becoming slightly massicot 
yellow or naphthaline yellow on throat. 

Male. Black above (only under lens are any stripes revealed, except stripe 
from snout through eye halfway to groin, which is black). Light stripe on 
upper jaw as in female. Throat yellow ocher or old gold. Venter white. Under- 
side of hind legs as m female. While in hand the upper parts changed to buffy 
olive or grayish olive. 

Structure: Vocal pouch round and subgular. 

"On the forefoot the second toe (counting from the outside) is much longer 
than the others, which are short; on the hind foot, very much longer still; webs 
of the hind feet take up not quite half the room between the toes; ins golden 
colored; a dark brown stripe runs from the nose through the eye up to the 
hind leg; the edge of the upper jaw white; all the upper part an inconspicuous, 
dark greenish gray, with three longitudinal rows of large, elongated spots of 
the same hue (only somewhat darker) on the back, . . . belly white; throat 
and chin brownish-olive colored; under side of the hind legs reddish gray. . . . 
Length i l //i length with hind legs extended 2'g"; length of the longest fore- 
toe 2 1 / 4"; length of the longest hind toe 4%" " (M. zu Wied, Gen., 1839, I, 
249-250). 

Voice: The call is a vibrating chirp. 

"During the last weeks of March Chorophilus (Pseudacris) appears in con- 
siderable numbers about the outskirts of Buffalo. The male chorus, which is 
easily distinguished from that of Hyla pic\enngii, rises from most of the 
swamps and temporary ponds, even within the city. The singers themselves, 



TREE FROGS: HYLIDAE 255 

however, are not easily seen, for, upon approach, they become silent and further 
disturbance causes them to disappear into the vegetation at the bottom of the 
pond where they remain until some time after the disturbance has ceased. 
Then, from some remote corner the chorus is gradually taken up until the 
whole pond resounds with the ringing notes. In taking up the chorus, the as- 
surance evidenced by the single voice is extremely contagious. This fact makes 
it possible to overcome some of the difficulties which ordinarily present them- 
selves in collecting this species. One has but to place the first captures in a bag 
or other close receptacle and carry it on one's person. The prisoners chirp up 
and sing as boldly as though undisturbed in their natural haunt. Their voices 
elicit an almost immediate response from those in the pond. Indeed, at such a 
time, with a little practice one can wade about, while they sing on all sides and 
dip up as many as one desires. In this way after spending several hours in cap- 
turing the first two, 30 specimens, 25 males and 5 females, were taken in less 
than an hour" (A. H. Wright and A. A. Allen, N.Y., 1908, pp. 39-40). 

"It has a note somewhat similar to the preceding species (Acns gryllus crept- 
tans), but the pitch is higher and the rattle is less definite. The note is seldom 
heard in daylight hours except on dark days. The writer has never heard it, 
as Cope &ays, in the hottest hours" (M. Morse, Ohio, 1904, p. 119). 

"With the first mild spring days, often before all the snow and ice of winter 
have disappeared, the loud trill of this small species may be heard from pools 
and ditches. The note is so resonant that on quiet evenings it may be heard a 
half mile or more and is commonly attributed to the larger frogs of the genus 
Rana. When the note is uttered the vocal sac is extended to its utmost and is 
larger than the head" (H. Garman, 111., 1892, p. 345). 

"The swamp tree frog is found in marshes and damp places throughout the 
summer and fall. During this time it is solitary and its call is rarely heard. It is 
also seldom seen because of its small M/C and protective coloration. When dis- 
turbed it disappears in the water, but it is a very poor swimmer and soon comes 
back to land. ... It comes from hibernation early. The song is very loud. 
When croaking, the male sits upright in the water, supporting himself with 
grass, leaves or twigs, and sings with the head and vocal pouch out of the water. 
When disturbed, he sits perfectly still and does not resume his song until 
the source of the alarm has passed" (A. G. Ruthven, C. Thompson, and H. 
Thompson, Mich., 1912, pp. 47-48). 

Breeding: This species breeds from March 20 to May 20. The egg mass is a 
loose irregular cluster, the mass small, less than i inch in diameter, 20-100 eggs 
in the mass, 500-1500 eggs in the complement. The brown or black and white 
egg is /^ri-%0 inch (0.9-1.2 mm.), the single envelope M-%o inch (5.0-7.8 
mm.). The tadpole is small, i: Ko inch (23 mm.), deep-bodied with a long 
tipped tail, quite black with bronze on the belly and sides, and the tooth ridges 
are %. After 40 to 90 days, the tadpoles transform in June at /io-%o inch (7.5- 
ii mm.). 



256 HANDBOOK OF FROGS AND TOADS 

"While travelling along the state highway in the southern part of Logan, 
Utah, on the i5th of May, 1919, I heard great numbers of the swamp tree 
frog, Pseudacris triscriata, uttering their characteristic songs. I found this am- 
phibian very common in small ponds at the roadside, and there were scores of 
egg masses, attached in most cases to blades of grass. No tadpoles had yet ap- 
peared. So numerous were the egg masses that I collected a representative lot 
of them. The number of eggs in the twenty-two egg masses taken were as fol- 
lows : 66, 45, 53, 33, 65, 46, 88, 38, 40, 67, 32, 50, 64, 87, 77, 15, 65, 51, 73, 45, 130, and 
190. The number of eggs here found is much greater than that typical of the 
species as given by Dickerson in The Frog Book, i.e., 5-20 (page 159). The 
masses containing 130 and 190 eggs respectively seem extreme. There is a possi- 
bility that two egg masses became fused in these cases, but except for the fact 
that the numbers of eggs are unusually high there is no reason for believing so, 
as the gelatinous material remained in one compact body, and gave no evidence 
of a multiple origin" (H. }. Pack, Utah, 1920, p. 7). 

"Eggs and adults of this little frog were taken in temporary pools formed by 
the melting snow along a railroad right-of-way near Boulder, during the first 
ten days of May, 1914. On May 9 eggs were collected in the four<ellcd stage 
and kept out of doors in water from the pool in which they were found. The 
development of these eggs was very rapid, a fact which may be correlated 
with the use of temporary pools as the spawning grounds by this species. On 
the nth all the eggs were in the elongated stage preceding hatching, and dur- 
ing the 1 2th most of the eggs hatched. The tadpoles of Chorophiltts trisenatus 
immediately after leaving the eggs were very black and about 8 mm. in length, 
resembling the tadpoles of the common toad in outline" (M. M. Ellis and 
J. Henderson, Colo., 1915, p. 257). 

"May 17, 1919, Fort Sheridan, 111. Found Pseudacris jenarum \trisenata\ 
breeding in trenches on the Rifle Range and collected a couple of egg clusters. 
The eggs are attached to grass stems arranged in a layer one or two eggs thick 
all around the stem, forming a mass about two inches long and less than half 
an inch in diameter. . . . June 22. Tadpoles of this species are metamorphosing 
in the same pools on the Rifle Range where eggs were found on May 17. They 
were in all stages of development, from tadpoles two-thirds grown, with short 
hind legs, to young frogs with tails nearly absorbed. The measurements of sev- 
eral specimens follow : Two full-grown tadpoles with short hind legs measured 
26 and 26.5 mm. Four with large hind legs and fore-legs almost ready to burst 
out; 25.5, 27, 27.5, and 28 mm. Four young frogs with all four legs well devel- 
oped and tails in different stages of absorption measured 10, 10, 10, and n mm. 
in body length" (P. H. Pope, 111., 1919, pp. 83-84). 

"Its eggs were found in a small pool on the 22d of March. They were at- 
tached to twigs in small and large bunches. Each egg was one-third inch in 
diameter, including the usual coating of jelly. . . . The tadpoles were set free 
on April 5. They are slenderer than the larvae of the Leopard Frog, and not so 



TREE FROGS: HYLIDAE 257 

dark in color. They arc dark gray rather than black. . . . The rudiments of 
the hinder limbs appear about the 2oth of April. At the same time two rows of 
horny teeth appear on each lip, and a few days later an additional row on the 
lower lip. These teeth are minutely denticulated at their tips, and they form 
an admirable apparatus for scraping off the layer of nutritious slime that covers 
all objects in the water. There are from 55-95 of these teeth in each row. . . . 
By April 20 the length has become one-half inch and by May 4 about three 
quarters. The body is of a dark color, adorned with numerous blotches of gold. 
The belly is nearly covered with a shimmer of gold and coppery. When three- 
fourths inch long the young were observed to come to the surface and take in 
air. By the 2oth of May the young have attained the total length of a little over 
an inch. By the 26th of May the tadpoles had attained a length, in some cases, 
of 28 mm., 16 of which is tail" (O. P. Hay, Ind., 1889, p. 773). 

"Many of them about this time succeed in releasing their forelegs from tfie 
skin which has held them down. Now the tadpoles grow smaller instead of 
larger. This is largely due to the shortening of the intestine at this period of 
transformation. These four-legged tadpoles are very lively and very timid. 
They show a great inclination to get out of the water and to hop about. They 
soon lose their skill in swimming, and if confined to the water too long will 
drown. The disks are seen on their fore feet as soon as these feet appear. The 
tails are rapidly absorbed, and by the i2th of June all have become little frogs 
like the adults, except in size, At the time of transformation the length of the 
head and body is less than one-half inch" (same, p. 471). 

"In Kansas the eggs arc more numerous in their respective masses than these 
previous accounts would indicate, numbers varying from 110-300, the mode 
being about 140. Wright and Wright [Gen., 1924, p. 381] give 500-800 as the 
egg complement. One female observed by the writer laid 1,459 eggs during a 
single night in the aquarium in which she was isolated. The membranes of 
the eggs from females near Lawrence differ somewhat from the type figured 
by Wright and Wright [Gen., 1924, pi. i, fig. 7], The outer envelope is typi- 
cally about 3.0 mm.; there was, in several bunches of eggs, a second membrane 
about 2.1 mm. in diameter; the vitellus is as given by Wright and Wright (0.9 
to 1.2 mm.). . . . There is always a vitelhne membrane closely applied to the 
vitellus" (H. M. Smith, Kans., 1934, PP- 4^7> 468). 

In 1936 K. A. Youngstrom and H. M. Smith (Kans., 1936, pp. 629-630) de- 
scribed the tadpoles of this species. The distinctive thing they showed is lower 
labial tooth rows 3 instead of 2. This is a fact that like many others we dis- 
covered soon after our 1929 paper. 

Journal notes: Ithaca, N.Y. Frogs from Buffalo, N.Y. On the morning of 
April 2, 1907, the 25 males were placed with the 5 females. At 9:50 A.M. the first 
pair was recorded and within 20 minutes the female began laying. With one 
exception, she chose a different perch for each egg-laying period, thus giving a 
bunch to each period of sexual activity. In one instance she sought the same 



258 HANDBOOK OF FROGS AND TOADS 

perch three successive times, This pair consumed 2% hours in laying a com- 
plement of 500-600 eggs. The process required about 90 fertilizations and emis- 
sions. Each time from two to ten eggs were voided, being emitted in small 
strings, a condition readily seen when occasionally the eggs were unattached 
and hung down from the vent of the female. All the eggs were laid in water 
66F. 

April 4, 1928, Hilton Beach, Hilton, N.Y. The common name, "Chorus 
frog," is very appropriate. Only a few toads, meadow frogs, and peepers were 
speaking. It was a very deafening chorus, not like Hyla femoralis and some 
larger Hylas but very pronounced. This Swamp Cricket Frog may be in 
swampy places where there is brush, but it can hardly be kept down in note to 
a cricket when in chorus. 

In New York I had never heard it except at Hilton, Hilton Beach, Buffalo, 
and Hamburg. By its chorus, we found it in Greece, April 4, at Hilton and 
Hilton Beach April 4 and 5, and at Buttonwood Creek at Parma and Greece 
town line. On April 5 heard it near Rochester, near Webster, Williamson, near 
Westbury, 3 miles north of Auburn on the Auburn Weedsport road. 

April, 1928, Hilton, N.Y. West of Mt. Reed Church, Greece, we found it m 
a pond in an orchard where several small trees and bushes were, and also brush. 
The frogs seemed to be in a grassy field, but mostly along a fence-row pond 
with trees and brush. Along Parma and Greece town line, they were in over- 
flow pools of Buttonwood Creek. Not brushy there. They were also in pools 
in grassy fields toward Braddock's Bay. 

May 30, 1942. With Wilmcr Webster Tanner went i mile west of Provo, 
Utah, to a spring. Here heard a great chorus of P. n. triseriata. Anna caught a 
female. 

June 8, Flagstaff, Ariz. Went to Museum of Northern Arizona. Met E. D. 
McKee, H. S. Colton, Major Brady. At museum saw Pseudacns n. tnsenattis 
from Apache County. . . . Tonight went to Mary's Lake, 7 miles long. At 
upper end a great chorus of P. n. tnsenatus. When we returned, in a stream 
back of the motel heard several P. n. triseriata. 

Authorities 9 corner: 

F. W. Cragin, Kans., 1876-81, p. 121. A. R. Cahn, 111., 1926, pp. 107-108. 
B. W. Evermann and H. W. Clark, F. G. Evenden, Jr., Ida., 1946, p. 257. 

Ind., 1920, pp. 635-636. 

Florida Chorus Frog, Florida Winter Frog 

Pseudacns nigrita verrucosa (Cope). Plate LIV; Map 17. 

Range: "Peninsular Florida. Known from the following counties: Alachua, 
Duval, Volusia, Marion, Putnam, Sumter, Lake, Hernando, Pinellas, Polk, 
St. Lucie, Charlotte, Lee, Collier, Broward and Dade" (Carr, Fla., 1940, p. 56). 

Habitat: Flatwoods, prairie lands, glade depressions, and ponds. "Swamps 



TREE FROGS: HYLIDAE 



259 




Plate LIU. Pseudacris nigrita triseriata Plate LIV. Pseudacris nigrita verrucow 

(XO- i*3i5/>- Females. 2. Male. 4. Eggs. (X0 l ~^> Males. 



260 HANDBOOK OF FROGS AND TOADS 

and grassy ponds and ditches; moderately common" (O. C. Van Hyning, Fla., 



Size: Adults %-i l /z inches (22-30 mm. or more). 

General appearance and structure: 'The length of the head to the posterior 
margin of the membranum tympani enters the total length to the vent three 
and one-sixth times. The head itself is narrow and acuminate, the muzzle pro- 
jecting acutely beyond the labial margin. The external nares mark two-fifths 
the distance from the end of the muzzle to the orbital border. The membranum 
tympani is only one-fourth the diameter of the orbit. The canthus rostrahs is 
distinct, but obtusely rounded. The vomerine fasciculi are approximated, and 
near the line of the posterior border of the nares, which are larger than the 
minute ostiu pharyngca. The tongue is large and wide behind and faintly 
emarginate. 

"The heel of the extended hind limb extends to between the orbit and nos- 
tril: the femur is short, while the tarsus is long, a little exceeding half the 
length of the tibia, and exceeding the length of the remainder of the foot, 
minus the longest toe. The skin of the gular and sternal region is smooth; of 
the abdomen, areolate. That of the dorsal region is tubercular, smooth warts of 
large and small size being irregularly crowded over its entire surface, and not 
at all resembling the areolate surface of the belly, 

"Color above leaden, with three longitudinal rows of darker, light-edged 
spots, extending, one on each side, and one on the median line. They are each 
composed of a series of spots joined end to end. Femur and tibia cross-barred. 
Upper lip dark plumbeous, with a series of five white spots; a similar spot be- 
low the tympanum. Inferior surfaces yellowish. . . . 

"From Volusia, Florida; Mrs. A. D. Lungren. 'This Chorophtlns is similar 
in proportions to the C. triseriatus, but is well distinguished by the characters 
of the skin, and the coloration. The tubercular upper surface is quite peculiar, 
and the smooth gular region is equally wanting to the Northern Frog. The 
dorsal skin is somewhat like that of Acns gryllus " (E. D. Cope, Fla., 1877, 

PP. 87-88). 

"Cope is clearly in error in stating |Gcn., 1889, p. 338] that verrucosus 
differs from C. nigritus (Le Conte) in the somewhat longer tarsus, and 
from C. jenarum (Baird) in the longer hind leg. He records the tarsus of 
nigritus as 9.4 mm., and the 'hind leg' of jeriamm as 1.77 in. (45 mm.), as com- 
pared with a 'hind limb 1 of 26 mm. in verrucosus. The type of verrucosus is 
unquestionably immature" (M. K. Brady and F. Harper, Fla., 1935, p. 108). 

These authors emphasize "the maxillary stripe broken to form a series of 
irregular dots and bars or lacking" (A. F. Carr, Jr., Fla., 1934, p. 32). 

Voice: "During the night of January 20, 1932, R. F. Deckert and I were col- 
lecting in the Royal Palm Hammock, on Paradise Key. In this tropical setting 
we were surprised to hear the trilling of Pseudacris coming from the 'glade' on 



TREE FROGS: HYIJDAE 261 

the west side of the hammock. The call, though possibly a little higher in pitch, 
was remarkably similar to that of P. n. feriarum as heard in the vicinity of 
Washington, D.C. The chorus was very small and localized. After considerable 
difficulty we managed to approach the position of the calls and found them 
coming from depressions and 'potholes' among the limestone rocks and 'glade* 
vegetation. These depressions contained shallow ponds that were choked with 
hnardia. Once we had gotten into their immediate vicinity, the frogs became 
silent, but by running a finger along the teeth of a pocket comb, we were able 
to imitate the voice sufficiently to induce them to call again. We thus located 
the males, which occupied well-separated stations and were sitting on projec- 
tions of the limestone above the water line of the little ponds" (M. K. Brady 
and F. Harper, Fla., 1935, p. 109). 

"The call is similar to that of nigrita, but the individual crepitations come 
much faster perhaps twice as fast and to some extent suggest the song of 
fmarum" (A. F. Carr, Jr., Fla., 1940, p. 56). 

Breeding and Journal notes: "Flat woods; prairie lands of the south-central 
peninsula. Not common. Habits apparently similar to those of nigrita. Breed- 
ing February to August 15, in cypress ponds, flooded meadows, drainage 
ditches in the muck land around Lake Okecchobce, and in 'pot-holes* and 
ditches in the South Florida limestone" (A. F. Carr, Jr., Fla., 1940, p. 56). 

On Feb. 7, 1932, O. C. Van Hyning of Gainesville, Fla., wrote as follows: 
"I went out into the middle of the Prairie a few nights ago, where there was 
a little water, and caught eight males and one female of l\ nigrita; the female 
and one male I watched from the time they were about a foot apart, until they 
mated, and would have stayed on another hour or so, but the mosquitoes had 
just about eaten me up, so I caught them. They have so far failed to mate again 
although two males were together once. The amplexation was axillary. I am 
going out to the same place as soon as I finish this letter, and see if I can get any 
additional females and see the eggs deposited." 

The following week, on Feb. 15, he wrote: "I have . . . two pairs mated, 
one of a male and a female, one of a female and two males, and one case of two 
males [in the laboratory]. I have one bunch of eggs laid in the laboratory, but 
they were pretty well walked over before I rescued them. I didn't know 
whether they were fertile or not and have preserved them." 

Brady at Royal Palm Hammock wrote: "In addition to a series of six calling 
males, we took a pair in amplexus. The female deposited 160 eggs, a few sep- 
arately, but the majority in a loose mass not differing from the egg mass of 
feriarum. The brown and white eggs resembled those of jeriamm, but seemed 
somewhat smaller, the vitellus measuring less than 0.5 mm. The single enve- 
lope measured 1.5-2 mm. Development was very rapid and hatching took place 
within 60 hours" (M. K. Brady and F. Harper, Fla., 1935, p. 109). 

On Feb. 4, 1933, at Gainesville, Fla., Messrs. A. F. Carr and H. K. Wallace 



262 HANDBOOK OF FROGS AND TOADS 

found this species breeding at the same time as P. ocularis and P. ornata. The 
eggs they sent us were brown and cream or white, the vitellus ^5 inch (0.9-1.0 
mm.), the single envelope Ko-% inch (2.6-2.8 mm.). 

Cope emphasized the warty upper surface. We have seen very warty ones 
in the southern half of the peninsula, and W. J. Hamilton, Jr., at Ft. Myers 
March n, 1940, took very tubercular specimens. 

Authorities 9 corner: 
A. F. Carr, Jr., Fla., 1940, p. 55. 

Western Chorus Frog 
Pseudacris occidentalis (Baird and Girard). Plate LV. 

This frog probably does not exist. 

If Cope (1889) was unwarranted in assigning Chorophilus occidentalis 
(B & G) (1853) to the eastern Rana ornata Holbrook (1836) or Cystignathus 
ornatus Holbrook (1842), what then is this Lttoria occidentals which Baird 
and Girard described in a paper entitled "A List of Reptiles Collected in Cali- 
fornia by Dr. John L. Le Conte . . ."? We prefer to believe it was collected 
in California. All the other batrachians of the report were strictly western 
forms. Baird and Girard report Hyla regilla in this same paper (Calif., 1853, 
p. 301), first describe it in another place of this same volume (p. 174), and Hal- 
lowell also describes Hyla regtlla in the same volume (p. 183) as Hyla scapu- 
laris. Nevertheless the Litona occidentalis of this report might well be another 
variant specimen of the very variable Hyla regilla. At that period we have Hal- 
lowell in the one paper (Calif., 1856, pp. 96-97) describing Hyla regilla three 
times as Hyla scapulans, Hyla nebulosa, and Hyla scapulans var. hypochondri* 
aca. We therefore suspect Litona occidentalis of being a synonym of a western 
Hyla, presumably Hyla regilla. A glance at the plate of Hyla regilla (Plate 
LXX) will show at once how many resemblances this species has to Pseudacris. 

In 1889 Cope (Gen., p. 337) speaks of C. occidentals (which we now term 
P. ornata) as ranging "to the Wichita River, in north central Texas. Specimens 
were sent me from the latter locality by that excellent naturalist, Jacob Boll, of 
Dallas. ... It does not occur in California as supposed when first described." 
What are these Texan forms? 

The records of three friends, C. S. Brimlcy, J. Hurter, and J. K. Streckcr, arc 
yet to be discussed. Brimley (Ala., 1910, p. n) secured "Chorophilus occidenta- 
Us from Bay St. Louis, Mississippi, February and April 1898, 5" and Choro- 
philus ornatus "Hastings, Florida, June, 1901, i; Green Cove Springs, Florida, 
July, 1898, 5." Doubtless a specimen in U.S. National Museum, no. 29189, la- 
beled C. occidentalis Hastings, Fla. H. H. & C. S. Brimley, is the one specimen 
Brimley rightly called C. ornatus above. He is the only person to receive each 
from the Southeast and the only one in that period to call the Floridan form 
C. ornatus. But what was the Chorophilus occidentalis of Bay St. Louis, Miss.? 



TREE FROGS: HYLIDAE 263 

In the Amphibians and Reptiles of Arkansas, Hurter and Strecker record 
Chorophilns occidentalis from "Hot Springs (Combs)." Strecker knows 
Pseudacris strecJ^eri (ornata of Cope, 1889) and has contributed as much as or 
more than anyone living to its habits, but the Pseudacris occidentalis is the 
puzzle. 

Since the paragraph above was written, there have come to hand Strecker's 
records of Pseudacris ornata (strecl^eri) and Pseudacris occidentalis. In recent 
times only two people have collected Pseudacris occidentalis in life to recog- 
nize it. Richard F. Deckert secured it in Jacksonville along with P. n. mgnta 
and P. octtlaris. This author does not record P. ornata and his descriptions of 
P. occidentalis are those of P. ornata, but the records of the other author who 
has recently seen this species cannot be so readily allocated to P. ornata (occi- 
dentals of Cope and Deckert), or P. strec\en (ornata of Cope and Strecker). 
The only collector in America who has taken P. occidentalis and the Texan 
P. ornata (now called P. strec\eri) in the same region, alive, is J. K. Strecker. 

On Sept. i, 1929, he wrote of the capture of specimens of each species in the 
Fort Worth region: 

"Western Chorus Frog (Pseudacris occidentalis (Baird and Girard)). I cap- 
tured two specimens. One, a junior, was caught four miles beyond Benbrook, 
in grass under the edge of a rock along a roadside; the second example, an 
adult, was caught among grass growing along a creek bank. Ornate Chorus 
Frog (Pseudacris ornata Holbrook). North and east of Fort Worth, I caught 
three adults among grass growing on the banks of small creeks" ( J. K. Strecker, 
Jr., Tex., 1929, p. n). 

In one museum some little P. octtlaris were termed P. occidentals. Of several 
different museum accessions considered P. occidentalis, we have made the 
positive note: "This is P. strecJ^eri. Possibly some of the Texan representatives 
may prove P. n. clarfyi" 

In 1930 we visited Dr. J. K. Strecker, Jr., and received many kindnesses from 
him. While we were in the field for P. strec\eri (ornata of Strecker), he re- 
marked that what he had recorded as P. occidentalis appeared different from 
jP. strecljfri (ornata) and had different habits. He kindly loaned us what speci- 
mens he had of P. occidentalis. At first they puzzled us, but the more we ex- 
amined them the more we inclined to interpret them as the spotted individuals 
of the so-called P. n. clarfyi. Taking his Nos. 4501 and 4502 (Baylor Museum) 
of Fort Worth and Bluff Creek, McLennan Co., and comparing them with his 
3507 P. nigrita, we call all three P. n. clar\ii. Another specimen, P. occidentals, 
from Gayle, La. (Frierson), may be a large female P. nigrita (triseriata) . 

Before we saw these specimens (4501, 4502) we wondered if they might be 
like P. strec\eri, but in longer head-to-tympanum, wider head, smaller eye, 
wider internasal, shorter hind limb, shorter first, second, third, fourth, and fifth 
toes, P. strec^eri does not overlap the relative measurements (number of times 
in length) of these two P. occidentalis specimens. 



264 HANDBOOK OF FROGS AND TOADS 

45 OI > 45 02 

Head to tympanum 3.16- 3.29 2.38 - 3.0 

Width of head 3.11-3.47 2.4 -2.88 

Internasal sp. 11.2-11.8 8.0 -10.2 

Eye 8.44- 9.33 9.33 -12.0 

Hind limb 0.66- 0.72 o-734~ 0-87 

First toe 5.45- 5.6 6.22 - 9.0 

Second toe 4.3 - 4.52 4.66 - 5.53 

Third toe 2.8 - 2.95 3.11 - 3.6 

Fourth toe 2.0 - 2.18 2.15 - 2.57 

Fifth toe 2.66- 2.68 2.8 - 3.5 

The relative measurements of Nos. 4501 and 4502 fall within the relative 
measurements of P. n. dartyi in almost every respect except head length and 
intertympanic and internasal widths. Most of the measurements of No. 1378, 
the specimen from Gayle, La., fall within those of P. n. triscnata. 

All in all our conclusions are that these P. occidentalis specimens may be 
forms of P. nigrita and are not P. streckfn (ornata of Strecker and Cope). 
Frogs called P. occidentalis in Georgia and Florida are P. ornata (Holbrook). 

Little Chorus Frog, Swamp Tree Frog, Swamp Cricket Frog, Tree Frog, 
Savanna Cricket (Bartram), Least Swamp Frog 

Psetidacns ocularis (Holbrook). Plate LVI; Map 18. 

Range: North Carolina to southern Florida. 

Habitat: Sphagnum edge of cypress pond. Grassy or sedgy area of pond or 
wet edge of pine barrens. 

May i8 3 1921, Pseudacris ocularis may start up from sphagnum edge of 
cypress pond or may start up from the ground. They will leap 1-1% feet at 
times. These specimens sat on Eriocaitlon, small Sarracema minor, sedge, and 
saw palmetto. We took a dozen of them. On June 25, we found them common 
in one pond, many in bamboo (smilax) vines, on bushes some 4-5 feet up, 
others on level of water. July 15, we found them abundant in outer edges of 
cypress ponds, margins of cypress bays. Most of them we caught in the grass 
near the edge of ponds. June 17, 1922, we found them in grassy and sedgy areas 
on wet edge of pine barrens or outer edge of cypress bay or branch. 

"Wet grassy places, especially in the pine flatwoods, common" (O. C. Van 
Hynmg,Fla., 1933, p. 4). 

Size: Adults, %-% inch. Males, 11.5-15.5 mm. Females, 12.0-17.5 mm. 

General appearance: These are the brownies of frogdom in the United 
States. They may be uniform gray, brown, greenish, or reddish on the back 
with a dark vitta from the eye backward as a stripe of variable length. This is 
set off by a light area below the eye extending backward to the shoulder. There 
may be a dark triangle between the eyes with a stripe extending down the 



TREE FROGS: HYLIDAE 



265 




iP ; - : ' ; fr "J- ^ : '-'"C;-^^ 
sis^^d^i^ 1 .^ 1 ! 

|iP^'>X--:-:- 



Plate LV. Pseudacris occidcntalis (X%)- 
1-7. Males. 8. Female. 



Plate LVI. Pseudacris ocularis. 1,4,5,7,8. 
Adults (XiVO- a/>- Males (Xi%)- 3- Male 
croaking (XO- 



266 HANDBOOK OF FROGS AND TOADS 

mid-back and a stripe on either side of this. These little frogs are so tiny, so 
delicate, that it does not seem possible that they are adult frogs. Their form is 
slender, their legs long, their eyes bright and beadlike, their snouts very pointed 
and extending beyond the lower jaw. The nostrils are on the sides of the 
pointed snout. These little midgets can turn their heads or tip them upward 
or sideways without turning the body. 

Color: On May 17, 1921, Okefinokee Swamp, Ga., we had diverse colorings 
in a lot of 16 in one pan. Some were tawny, hazel, or amber brown on upper 
surfaces; others buckthorn brown; one a light grayish olive or light olive-gray; 
another deep olive-gray. The stripe on the upper jaw was almost cream, chal- 
cedony yellow, or glass green. Belly chamois or cream-buff. Iris cinnamon. 

On Aug. 1 8, 1922, at Milliard, Fla., found a little Psettdacris ocularts which 
is cinnamon or ochraceous-tawny in color. It is an adult male with a plaited 
throat. The ten specimens vary in color from gray, greenish, light yellowish 
olive or dull citrine through to chestnut or bay; four specimens were of the 
yellowish olive or gray group, four of the chestnut or bay group, and two in- 
termediate. The most distinctive, yet not always constant, mark is a line of 
dark color which runs from the snout or nostril through the eye and tympanum 
to or beyond the arm insertion. In three specimens it runs halfway to the hind 
limbs. In two more it just passes the arm insertion, as Holbrook figures it. In 
two this dark stripe is absent. In the striped individuals the limbs bear cross- 
bars. In all material, the entire upper surfaces have a very fine speckling of 
close-set dark dots. The under parts arc whitish or yellowish white with the 
fine speckled dots of the upper parts all over the ventral side. In some the un- 
derside of limbs is almost as dark as the upper parts. 

When four specimens arrived May 7, 1929, from Gainesville, Fla., from O. C. 
Van Hyning, in sphagnum, all were different: one was uniform carnelian red 
above with no stripes except vitta on side; another dark olive; one deep olive- 
buff with three stripes on back; and one avellaneous. Later the carnelian red 
was entirely changed. 

Structure: Midgets in size; snout pointed and projecting beyond the lower 
jaw; nostrils on sides of snout; slender in form, hind legs very long; eyes bright 
and beadlike; disks on fingers and toes small but distinct; skin of back covered 
with very fine warts. 

"H. ocularis. H. ocularis Daudin, i c. p. 32. Hyloclcs ocularts Holbrook, pi. 
35. Above brown or bronzed or silvery grey, very finely speckled with dusky 
or darker, a tolerably wide band of black proceeds from the tip of the nose to 
the middle or beyond the middle of the sides, this is bordered beneath with 
white. Chin and underside of the thighs speckled with black. Legs speckled 
like the back and more or less spotted and barred with dusky, fingers and toes 
all furnished with small disks. Length .6. Inhabits Georgia. The smallest of all 
known Ranina, From the small size of this and the preceding species, the web 



TREE FROGS: HYLIDAE 



267 



between the third and fourth toes is not very perceptible" (J. Le Conte, Gen., 
1855, p. 429). 

Voice: The call is a high, shrill, cricketlike chirp or trill. Vocal sac of the 
male even when collapsed sometimes appears to cover almost half of the venter 




/VX Hyh regilla 

. wright or um 

Pseud acris ocularis 

P. n. clarkii 
I". P. n. sepfenfr/ona//s 

P. n. triseriata 



Map 18 

of the body. The rear of it may reach caudad to the pectoral region, to the line 
connecting one arm insertion with the other. Our first acquaintance with a 
chorus came the evening of May 20, 1921, in Okefinokee Swamp, Ga. Went 
down to the pond east of the Negro quarters and it sounded as if bedlam had 
broken loose. I heard a cricketlike note everywhere. It was Pseudacris ocularis. 
One frog was on a grassy mat, one on a log, another calling from pine brush 
at the edge of the water, another on the bole of a tree. They do not like the 
electric spotlight. One on pine brush worked up into the leaves. We flashed one 



2&I HANDBOOK OF FROGS AND TOADS 

on the side of a black gum. Its throat pouch was transparent and we could see 
through it and discern the bark behind. The croaks were given 30-65 per min- 
ute. The note is high-pitched, penetrating, can be heard 150-200 feet away. 
Whether characterized as chirp, trill, cheep, squeak, or high shrill insect call, 
it surely is a loud piercing call for so little a mite of frog flesh. It is an amusing 
little creature as it squeezes its slender body and throws out its large sac one- 
half the size of the body. 

Breeding: It breeds from January to September. The brown and cream eggs 
are single, laid on the bottom of ponds and in vegetation in shallow water, 
about 100 in number. The egg is Mo-Ho inch (0.6-0.8 mm.), the single envel- 
ope ^o-Via inch (1.2-2.0 mm.). The greenish tadpole is small, ! % inch (23 
mm."), its tail long. Its tooth ridges are %. After 45 to 70 days the tadpoles trans- 
form, June 30 to August 18, at '}io-% inch (7-9 mm.). 

Journal notes: In 1921, on July 3, we took a handcar trip to Honey and Black 
Jack Islands. It was a misty day at first, then sunny a while; it rained all the 
afternoon, however. The temperature was from 71 -76. On the sphagna- 
ceous strand of Honey Island heard these creatures near Black Jack Island; in 
mid-afternoon on Honey Island, in Cypress Bay between the Pocket and Jones 
Island, and all over the prairie, in sphagnum of wet thickets especially. Also all 
along the trestle between Jones Island and Billy's Island. Never heard such a 
frog din. 

On July 29, 1921, we were caught in a downpour on Black Jack Island. We 
were encamped in the moist pine barrens. All about was the strongest chorus 
of Psettdacris ocularis we ever heard. Temperature about 75. 

In 1922, we secured them on Chesser Island, near old Suwannee canal, along 
the road to Folkston, in ponds near Trader's Hill, near an overgrown sphag- 
num bog outside the swamp, and along the road from Folkston, Ga., to Jack- 
sonville, Fla. 

Authorities' corner: 

W. Bartram, Gen., 1791, p. 278. J. E. Holbrook, Gen., 1842, p. 138. 

F. M. Daudin, Gen., 1801-1803, VIII, G. K. Noble, Gen., 1923, pp. 1-5. 

68-69. F. Harper, Ga., 1939, p. 148. 

Ornate Chorus Frog, Ornate Tree Frog, Ornate Swamp Frog, Swamp Cricket 
Frog, Ornate Winter Frog, Holbrook Chorus Frog 

Pseudacris ornata (Holbrook). Plate LVII; Map 16. 

Range: North Carolina (B. B. Brandt, 1933) to Georgia, Florida, and pos- 
sibly west along the Gulf to Louisiana. Alabama (Viosca). 

Habitat: Holbrook wrote of its habits: "I have always found it on land and 
in dry places, and frequently in cornfields after light summer showers. It is 
very lively and active, making immense leaps when pursued, and consequently 
is taken with great difficulty." "The structure of this species indicates terres- 



TREE FROGS: HYLIDAE 269 

trial, possibly subterraneous habits. I have dug specimens out of the sweet po- 
tato hills in my garden" (R. F. Deckert, Fla., 1915). M. J. Allen found it 
breeding in grass-land pools. "Grassy ditches and ponds; moderately common" 
(O. C. Van Hyning, Fla., 1933, p. 4). 

Size: Adults, 1-1% inches. Males, 25-35 mm - Females, 28-36 mm. 

General appearance: This is a large Pseudacris, looking almost like a small 
Rana sylvatica. The general color of this frog is chestnut brown with two in- 
distinct darker dorsal bars and an indistinct darker spot between the eyes. 
There is a prominent dark, almost black, vitta from tip of snout through eye 
over tympanum, usually beyond the arm insertion, often extending halfway to 
the groin. Two frogs with more prominent stripes have, between the eyes, 
Y-shaped triangles, which in the other four do not show. The groin spots are 
inclined to be almost black oblique bars, with light outlines. In fact, the spots 
on groin are on a yellow background. The tiniest rim of the upper jaw is dark, 
but above this is a conspicuous light line extending to the arm insertion and 
broadest under the eye. On the front of the upper arm is a dark bar. The eyes 
are very prominent with the upper part of the iris a bright gold band. The 
throat of the male is dark olive, with a central longitudinal plait. The throat of 
the female is light. The legs are barred. 

Cope characterized it well enough but added to the confusion by calling it 
Chorophtlus occidentalis B. & G. 

"Muzzle rounded in profile, projecting. Skin of upper surface smooth. . . . 
More slender [than C. ornatus\\ width of head entering length 3 to 3.5 times; 
nostril nearer end of muzzle than orbit; posterior foot longer, not webbed, and 
without subarticular tubercles; heel reaching middle of orbit" (E, D. Cope, 
Gen., 1 889, p. 332). 

G. A. Boulenger, who called it Chorophtlus copti, wrote: "Vomerine teeth 
behind the level of the choanae; tibio-tarsal articulation not reaching the eye; 
back immaculate" (Gen., 1882, p. 332). 

ft Chorophtlus copii. Tongue circular, entire vomerine teeth in two slightly 
oblique oval groups behind the level of the choanae. Head slightly longer than 
broad; snout rounded, as long as the diameter of the orbit; canthus rostralis 
rather indistinct; interorbital space as broad as the upper eyelid; tympanum 
somewhat more than half the diameter of the eye. Fingers and toes moderate, 
the tips very indistinctly dilated; first finger shorter than second; toes with a 
slight rudiment of web; subarticular tubercles small; no outer metatarsal tu- 
bercle. The hind limb being carried forwards along the body, the tibio-tarsal 
articulation marks the tympanum. Skin smooth above, granulate on the belly. 
Olive above; sides with three black spots, viz, a streak from the eye to the 
shoulder, an oval spot in the middle of the side, and one or two smaller ones 
on the loan. Georgia. A 9 Georgia. 

"Mr. Cope considers this frog specifically distinct from C. ornatus and iden- 
tifies it with Hyla ocularis of Daudin. The latter opinion I cannot share, as 



270 HANDBOOK OF FROGS AND TOADS 

nothing in the French author's description and figure indicates the least anal- 
ogy with this form. The toes are represented as half-webbed in H. ocularis 
and the digital expansions of C. copii are so slight that they would certainly 
have escaped Daudin's observation. Moreover the colour is quite different" 
(same, p. 334). 

Color: Gainesville, Fla., from O. C. Van Hyning, March 24, 1932. First male. 
The background color of the back and top of head is sayal brown or tawny- 
olive, becoming on top of fore limb and eyelids and top of head cinnamon or 
orange-cinnamon. The hind legs may be verona brown. The band on either 
side of middle of back is indistinct and is snuff brown or Saccardo's umber. 
The stripe from snout obliquely upward to nostril, thence through eye over 
tympanum, shoulder insertion, two-thirds of way to groin, may be black or 
sepia in front and warm sepia at the caudal end. The one or two groin spots 
may be warm sepia, chocolate, or walnut brown. The three bars on femur are 
the same color. The interstices of groin spots are light cadmium to lemon yel- 
low. There are also flecks of the same on the front of the upper half of the fe- 
mur. The front of the femur is partly dark vinaceous-brown. The rear of femur 
has punctulations of vinaceous-brown or dark vinaceous-brown. Lower end of 
rear of femur has a few spots of lemon yellow or lemon chrome. The side of 
body is vinaceous-buflf, at times becoming on top of hind legs and side light 
drab or pale drab-gray. The light band on upper edge of upper jaw is pale dull 
green-yellow, becoming at angle of mouth and under the tympanum white. 
The dark vitta from eye backwards on its upper margin has a thin outline of 
seafoam yellow to white; the groin spot or spots have a suggestion of the same 
thing. The upper jaw rim is a thin line of buffy brown. The eye color in the 
lower half partakes of color of the vitta, wherein the upper half is the color of 
top of head. There is a prominent natal brown or vinaceous-brown band from 
the lateral vitta down across the arm insertion extending even onto the forearm 
to make its first crossbar. On forearm, there is a prominent bar of same color 
at base of hand. The underside of fore and hind limbs is brownish vinaceous. 
The belly is a dirty white with considerable spotting in the rear pectoral region 
of drab or olive-brown. The throat is pronouncedly discolored, its rear sharply 
outlined by a fold; the caudal half is olive-ocher or old gold, the cephalic por- 
tion is Saccardo's olive, the two central plaits becoming olive-citrine. The edge 
of the lower jaw is spotted with white or pale dull green-yellow. After we had 
finished describing it, the dorsal color came to be uniform with no suggestion 
of long dorsal stripes. 

Second male. A uniform mahogany red, chestnut, or bay on the back. 

Female. Charleston, S.C., from E. B. Chamberlain, Nov. 4, 1932. The back- 
ground color of the back and top of head is Hay's russet, becoming on side 
orange-cinnamon or sayal brown. The two parallel dorsal bands are liver 
brown or carob brown, so also are the bars on the femur and tibia. Those on 
tarsus and foot merge into black. Snout stripe, eye vitta, lateral and groin spots 



TREE FROGS: HYLIDAE 271 

are light cadmium or lemon yellow. The rear of the femur much as in the male. 
The light band along the upper jaw is very narrow and whitish. The upper jaw 
rim is bone brown. The eye is like that of the male. The venter is more or less 
like that of the male, except that the throat is white without discolor. 

Structure: Form more slender than Pseud acris strecf^eri; head narrower than 
P. stred(eri; arms and legs longer and more slender than P. strecl^eri; fingers 
and toes long and slender, with mere trace of web at base of toes; snout pointed ; 
nostril equidistant between snout and eye; vomerine teeth between nares. 

"The head is small, with a broad, indistinct, triangular spot between the or- 
bits, the apex of which is directed backwards. A black line extends from the 
snout to the orbit of the eye, including the nostrils; below this black line is a 
yellowish blotch, covering most of the upper jaw. The lower jaw is cinereous 
above and white below. The mouth is small, and the palate is armed with two 
groups of exceedingly minute teeth between the posterior nares. The nostrils 
are placed on a slight prominence. The eyes are large and projecting, the pupil 
very dark, the ins of a golden colour. The tympanum is small, very dark col- 
ored, and placed in a dark vitta, or blotch, which extends behind the orbit to 
within a short distance of the shoulder. The body is short, of a delicate dove- 
colour above, with two or more oblong spots of dark brown, margined with 
yellow, on each side of the vertebral line; below these, and on each flank, are 
three smaller spots, likewise margined with bright yellow, the anterior one 
being the largest; these, with a smaller one above the vent, form a triangle on 
each flank. . . . The toes are five in number, not palmated, the two outer ones 
only are united at the base. Dimensions. Length of body from the snout to the 
vent, 1% inches; of the thighs, % an inch; of the leg, % an inch; of the tar- 
sus and toes, nearly % of an inch" (J. E. Holbrook, Gen., 1842, pp. 103-104). 

Voice: "This call is very loud, similar in pitch to that of Hyla picl(enngi, but 
much shorter, and at a distance sounds like the ring of a steel chisel, when 
struck by a hammer . . ." (R. F. Deckert, Fla., 1915). 

Norman Davis of Gainesville, Fla., says, "The call reminds me of Hyla cru- 
cifer but is without the trill." 

"I have timed ornata and found the calls to be repeated from 65-80 times a 
minute. I have never actually timed crucifer, but its calls are a good deal slower, 
probably 40-50 per minute. Ornata 's call is a single sharply terminated note, 
while crucifer s is a more deliberate slur from a lower to a higher" (letter from 
A.F.Carr,Feb.5,i933). 

Breeding: Winter months. November, December, January to March depend- 
ing upon the rains. H. K. Wallace and A. F. Carr, Jr., at Gainesville, Fla., se- 
cured eggs laid Feb. 5, 1933. They are brown and cream or white, and measure: 
the vitellus Vzs inch (0.9-1.0 mm.), the single loose envelope /4-% inch (3.6- 
4.2 mm.). They transform at % inch (14-16 mm.). Tadpole reddish brown, 
23-25 mm. 

Our evidence on transformation comes from two lots of material (USNM 



HANDBOOK OF FROGS AND TOADS 

and AMNH). The first accessions (USNM nos. 71030-3) were collected at 
Gainesville, Fla., by Gerrit S. Miller, Jr., and C. R. Ashemeier. They measure 
respectively 14, 15.5, 15, 14 mm. These specimens reveal none of the character- 
istic spots of the adult except the bars on the fore and hind legs. All four have 
an indication of the dark band from eye to shoulder. 

The other notes were taken on 19 "Pseudacris copit" specimens collected by 
T. Hallman in Florida (AMNH nos. 15155-8, 15234-47, 15976-7). My notes 
designate them as transformed specimens, but evidently some of them are 
slightly or well past transformation. 

They range from 14-19 mm. in length. Of two we write : No. 15157 18 mm., 
tail 9 mm., musculature light below, dark above. No. 15158 16 mm., tail 16 
mm., very spotty musculature, dorsal and ventral crests. 

The measurements of all were: 14, 15.5, 15.5, 16, 16, 16, 16.5, 16.5, 17, 17.5, 18, 
18, 18, 19, 19, 19, 19, 19, 19 mm. In 1922 Hallinan also took at Arlington, Fla., 
a transformed specimen (AMNH no. 215360). It is just transformed and 17 
mm. long. 

Finally three specimens (Fla. State Mus. no. 47679) taken 6 miles southwest 
of Gainesville, Fla., May 7, 1930, by O. C. Van Hyning and Norman W. Davis 
measure 16, 18, and 20 mm., respectively. The first is just transformed and the 
others, though past, still show signs of transformation. The transformed one 
of 16 mm. has a dark bar from eye to shoulder and a spot halfway from fore 
limb to hind limb. The 20-mm. specimen has the vitta from snout through eye 
and onto sides, also oblique groin spot and oblique pair of spots on back from 
anus forward. 

We still need detailed descriptions of the egg mass, individual eggs, and tad- 
poles. 

Authorities 9 corner: 
F. Harper, Ga., 1937, pp. 260-272. 
A. F. Carr, Fla., 1940^ pp. 57-58. 

[Note: See comparison with P. strec\en in the account of that species.] 

Strecker's Ornate Chorus Frog, Texas Ornate Chorus Frog 

Pseudacris strec\eri Wright and Wright. Plate LVIII; Map 16. 

Range: Texas, Oklahoma, southeastern Arkansas. The Mississippi record 
may have been a mistake. Was it P. brachyphona or P. ornata? Bragg (1942) 
records it from 19 or 21 counties in Oklahoma, and we have seen it from Bee- 
ville to Waco in Texas. 

Habitat: Moist shady woods, grassy pastures, among grain stalks, or in cot- 
ton fields. We found it breeding in a semiswampy, springy fork of a rocky 
branch of Helotes Creek in Texas. From our first experience with it and Streck- 
er's site (1910) we concluded it preferred rocky ravines or deep gulches, but 
the males are not choosy at all. 



TREE FROGS: HYLIDAE 



273 




Plate LV1I. Pseudacris ornata (X')- i-4- 1>late LVUL ^udacris streckfri. 1,3,4,5,6. 

Females. Males (XO- 2- Male (X%). 



274 HANDBOOK OF FROGS AND TOADS 

Strecker writes (Tex., 19266, p. n) : "These frogs inhabit all kinds of situa- 
tions. We found them in moist shady woods, in weed clumps and among grain 
stalks along the edges of corn, wheat, oat, and cotton fields, among weeds 
along the sides of dry, dusty roads, in grassy pasture land near lagoons and 
streams, and among sparse vegetation on the crests of low rocky bluffs. They 
were lively and seemed perfectly comfortable under the hot rays of the sun. 
Several specimens flushed by us sprang into small bushes and clung to their 
branches." 

Size: Adults, 1-1% inches. Males, 25-41 mm. Females, 32-46 mm., possibly 
ripening as early as 27 or 28 mm. In Strecker's collection there is not much 
evidence of ripening at 28 mm. 

General appearance: The head is short and wide; the body, short and 
squatty. The fingers of the hand are chubby, forming a toadlike hand. The 
toes are very slightly webbed, the disks slight. The frog varies in color from 
pale drab-gray or pale smoke gray to hazel, brownish olive, or pea green. 
Usually there are dorsal spots and frequently a triangle between the eyes. A 
dark line extends from the snout through the nostril and eye, over the tympa- 
num to the arm insertion; it is sometimes continued as spots along the side. 
Dark crossbands are on the legs. The groin and the front and rear of the femur 
in males may be yellow or green or olive-buff. The throat of the male is dark. 
The tympanum is very small. 

Color: Male. Helotes, Tex., Feb. 16, i()25. Back drab-gray; triangular spot 
between eye and spots on arms, legs, and back grayish olive; line on nostril, 
spot below eye, vitta back of eye, shoulder spot, and spots in a row along the 
side, black, and tympanum included in this color; crossbands on femur gray- 
ish olive behind, blackish in front; a black spot at the insertion of arm; grape 
green or dark olive-buff in groin and on front of femur; throat very discolored, 
dark olive-buff at its caudal edge shading forward into deep olive; under parts 
white; underside of thighs slightly dark olive-buff and buffy olive. Iris vina- 
ceous-fawn or light cinnamon-drab with "bronzy cast." There is a tendency for 
a horizontal line of black across the ins to connect the nostril line with the 
spot back of the eye. Another specimen brownish olive above with no dorsal 
spots. Another one with dorsal spots of calla or parrot green. In one, the spots 
through nostril, back of eye, over shoulder, and back of shoulder seal brown. 
In this specimen, groin, front and rear of femur, and tibia are lemon, chrome, 
or aniline yellow. Light color on upper jaw and rim of lower jaw back to un- 
der tympanum is olive-buff. 

Strecker (i<)26e) says, "The following are the three principal types of perma- 
nent marking: 

"A. Large and distinct black-brown dorsal markings in regular pattern. 
(Mostly males.) 

B. Distinct markings along both sides of the dorsal area, but absent from the 
middle of the back. (Mostly females.) 



TREE FROGS: HYLIDAE 275 

C. Markings of the dorsal region broken up into spots and flecks. A very dis- 
tinct and uncommon type, confined to a few female specimens." 

Of a series of 20 males and 12 females, Strecker (Tex., ig26d, p. 9) found a 
range of color thus : Light dove gray, dark gray, light plumbeous wood-brown, 
purplish brown, chestnut, bronze-buff, dull green, pea green, suffused with 
green above. He specifies the markings in each case. 

Structure: Tongue rounded behind. Vomerinc teeth behind nares. Very lit- 
tle webbing on feet. When wiping off one male for photographing, I sensed 
that it had considerable secretion. Very pebbly or granular on femur and ven- 
ter to pectoral fold. Two metatarsal tubercles. 

In preserved specimens a median plait on throat, or two plaits, or as many as 
six or more. Each side of throat often pebbled. The male sometimes, like the 
female, may have a transverse fold across the pectoral region. Some females 
may also have one across the base of the throat (in preserved specimens). 

E. D. Cope's first description of present P. st*cc\en follows: "Chorophihts 
oculans Daudm, Cope. This species resembles the eastern C. oculans f but some 
specimens differ in the tuberculate character of the skin of the superior sur- 
faces, and in the rudimental web of the hind foot. The head is rather short, and 
the anterior outline is a narrow oval. The extremity of the muzzle projects be- 
yond the mouth, and the lores are slightly oblique and a little concave. The 
nostril is but little nearer the extremity of the muzzle than the orbit. The verti- 
cal diameter of the tympanum a little exceeds the transverse, which is one-half 
the long diameter of the eye-slit. The pupil, as in the other species of this genus, 
is horizontal. The tongue is wide, discoid, and entire behind. The ostia pharyn- 
gea are smaller than the small choanae. The vnmerinc patches are short and 
transverse; they are entirely within the lines of the inner borders of the choanae 
and behind the line of the posterior borders of the same. 

"The tubercles of the superior surfaces are small and rather closely placed; 
they are largest on sides of the back. There is a faint arcolntum of the gulnr 
region. The limbs arc short and stout. The humerus is half or more inclosed in 
the skin. The palm reaches nearly to the end of the muzzle. The fingers are 
short and stout, and have neither dilations nor borders. The first is shorter than 
the second, which equals the fourth. The palmar tubercles arc not distinct. The 
heel of the appressed hind foot in thin specimens marks the middle of the tym- 
panic disc or posterior border of orbit, and the end of the muzzle the extremity 
of the tarsus. The hind foot beyond the tarsus is only as long as the tibia. The 
toes have no dilatations, but possess dermal margins, and a short but distinct 
basal web. There is but one solar tubercle, a small cuneiform prominence. 
Total length, m., .035; of head, to line of posterior borders of membramtm 
tympam, .011; width of head at the latter, .014; length of hind leg, .045; of 
femur, .013; of hind foot, .022; of tarsus, .CXJQ. 

"The color above is olive-gray, and below, uniform straw-color. A black band 
passes from the end of the muzzle on each side, through the eye, and expand- 



276 HANDBOOK OF FROGS AND TOADS 

ing over the ear-drum, terminates in front of the humerus. One or two dark 
spots above and behind the axilla may unite to form part of a lateral band. 
There may or may not be blackish spots above the groin and on the coccygeal 
region and anterior part of the back. The limbs have a few dark-brown cross- 
bands; the femur is yellowish and unspotted behind. 

"There is some difference between the Texas specimens and those from 
Georgia. Specimens from the latter State are very smooth, and the limbs, espe- 
cially the feet, are slender. The heel reaches to the orbit, or at least to the front 
of the tympanic membrane, and the end of the tarsus extends to or well beyond 
the end of the muzzle. The web and digital dermal borders are much less 
marked. Two specimens were obtained by Mr. Boll near Dallas, and three at 
Helotes by Mr. Marnock. All the latter have large brown dorsal spots" (E. D. 
Cope, Tex., i88oa, pp. 27-28). 

Cope compared this Texas form with P. ornata, which he called both C. ocu- 
lans and C. occidentalis. Workers now have a simple time in comparison with 
our struggles when we tried 30 to 40 years ago to identify Chorophtlus now 
Pscudacns. From 1900 to 1915 everyone was hopelessly confused in terminology 
and records. Field experience, of which we yet need much more, was the main 
factor in clarification. 

Voice: This call is rather shrill, somewhat like that of the common tree toad. 

"The calling habits of P. strecferi are distinctive. The voice is clear and bell- 
like and has considerable carrying power. The breeding cry is a single note but, 
when heard in a chorus, the sound resembles the squeak of an ungreased 
wooden wheel. This effect results from a lack of synchrony of individual calls. 
Males call from various positions. I have found them hanging to vegetation by 
their hands in water two feet deep, in vegetation above the water, and on the 
banks of pools, sometimes as far as five or six feet from water. One was found 
sitting on a floating board in a deep ditch calling lustily. In the more than fifty 
cases lhat I have observed, P. strecfon did not stand 'on tip-toe' while calling 
as does its close relative, P. ornata. . . . 

"The call of an individual of this species stimulates other males to call also. 
I have often approached a pool when no males were calling. Within a few 
minutes, when one call was heard, a whole chorus would break out and con- 
tinue lustily for ten minutes or more. Then all lapsed into silence again. After 
an interval of five to fifteen minutes this process would be repeated. This phe- 
nomenon is more prevalent during daylight than at night. Usually after dark- 
ness has fallen, especially after heavy rains within the breeding season, calling 
is continuous on through the night' 1 (A. N. Bragg, Okla., 1942^1, p. 49). 

Breeding: These frogs breed from December to late May. Our Texas friends, 
Mr. Kirn and Mr. and Mrs. R. D. Quillin, hold that they are more or less active 
and breed through the winter from December to late February. Mr. Garni at 
Bocrne, Tex., says, "They are heard throughout the winter, especially after 
rains." 



TREE FROGS: HYLIDAE 277 

"The eggs, like those of Pseudacris triscnata (Wied) are in small bunches, 
usually from ten to twenty-five. In the ditches leading to Belle Mead (Dry 
Pond), they are attached to weeds and water grasses, occasionally to twigs and 
water-logged branches" (J. K. Strecker, Jr., Tex., 19266, p. n). 

One egg mass in Strecker's collection is very large. There are approximately 
200 or 250 eggs in one loose mass. Each egg is brown above and cream below. 
Apparently, according to a superficial examination, there is one envelope, 
which is about 2% or 3 times the diameter of the vitellus, which is roughly 
2 mm. or less. Detailed measurements of these eggs reveal the following: 

The egg mass is rather milky in its formaldehyde preservation and has some 
precipitate on it. There seems to be only one envelope, though at times appear- 
ances indicate a second inner one; but of this we are not certain. Our impres- 
sion is one envelope, which measures from 3.0 through 3.6, 3.8 to 4.4 or 5.0 mm. 
in diameter. The vitellme diameter is from 1.2-1.8 mm., average 1.4, mode 1.4 
mm. 

"The tadpoles are small, blackish-brown creatures similar to those of Insert- 
ata, but are rather shorter and more robust. . . . About sixty days are required 
for them to go through with their complete transformation after leaving the 
egg" (J. K. Strecker, Jr., Tex., 19266, p. n). 

"The young specimens captured at Dry Pond near Waco, and at San Marcos, 
were about twelve millimeters in length, and must have been out of the water 
for several days, for the dorsal pattern was clear and distinct. When they first 
come to land, they are dark gray with hardly any trace of markings" (same). 

We have examined the Baylor material on which the previous statement is 
partially based. There are frogs, apparently just beyond transformation, meas- 
uring from 12 mm. (Baylor Mus. no. 3512) through 13 (3509) to 1^.5 (5228) 
mm. The body of the largest tadpole Strecker has is 12 mm. in length. 

The most precise information on the eggs and tadpoles comes from A. N. 
Bragg (Okla., 1942^ pp. 50-59), who has written: 

"The eggs are attached to vegetation in masses of from ten to about one 
hundred, just under the surface of the water. The usual number in a mass is 
from twenty to fifty and adjacent masses are often loosely joined by thin sheets 
of jelly. . . , The egg complement is of medium size. Three females produced 
the following numbers of eggs in the laboratory: 708, 695, and 401 and counts 
of several sets of adjacent masses of eggs collected in the field and presumably 
coming in each case from one female gave comparable values. More superficial 
observation of about fifty sets offered evidence that the above numbers repre- 
sent fairly closely the usual yearly egg capacity of one female. . . . 

"The eggs may be described as follows: color, brownish gray at the animal 
pole, the pigment extending downward to beyond the equator, then fading 
rapidly through shades of lighter gray to cream or nearly pure white at the 
vegetal pole. The animal pole is really quite dark and the color is only slightly 
less intense at the equator. The sizes of the eggs are nearly uniform and there 



278 HANDBOOK OF FROGS AND TOADS 

are no differences observable to the naked eye as one sees them en masse in 
culture dishes. Measurements of twenty live eggs by means of an ocular mi- 
crometer were as follows: 13 measured 1.23 mm. each; 4, 1.29 mm.; 2, 1.26 mm.; 
and i, 1.17 mm. . . . 

"The jelly is very sticky, almost absolutely hyaline, soft, and very elastic. 
Even in clear water, tiny particles of soil and algae catch easily upon it and 
serve to protect the eggs from detection. In a muddy pool, the jelly collects a 
complete coating of particles within a half hour so that its color becomes exactly 
that of the surrounding water. 

"The sizes of the individual masses of jelly of course vary with the number 
of eggs contained in each. The portion of jelly between each two adjacent eggs 
is also variable in amount but the distance separating each egg from the near- 
est one adjacent is usually between 5,58 and 7.25 mm. Staining of the jelly for 
one hour and forty-five minutes in Schneider's aceto-carmme revealed that 
(i) only one jelly envelope occurs about each egg; (2) this envelope is finely 
laminated, the jelly apparently laid upon the egg in concentric spheres, the wall 
of each of which is thin; and (3) there is a very thin, nongelatinous membrane 
surrounding a very small cavity in which the egg actually lies." 

Journal notes: Feb. 16, 1925, Hclotes, Tex. Anna says this is a fat squatty 
Hyla f not a Pseudacns. If is not in a class with Pseudacris oculans or Pseud- 
acris n. clarion. 

Feb. 16. Rather shrill call. At once it appealed as a note new to me. A note 
somewhat like H. verstcolor. Soon found three males calling. One was on a 
bank, another among vegetation in water 3 or 4 inches deep, and another hang- 
ing to a branch at the water. Put these in a bag, and they began to croak; had 
three or four diverse notes in the bag. These "bag frogs" made others croak. 
Caught five or six in all. Do not believe the species is at its height. Air 31 at 
7:30 P.M. when these were calling, water about the same. This afternoon was 
very humid. 

Feb. 19, Helotes, Tex. During the night, one or two captive males called in 
the can. Temperature close to ^2F. This morning air in the can is 41. Most 
of the frogs are rather inactive. 

Feb. 21. At 9 P.M. air 70, didn't hear them. When I returned, air about same; 
heard one, then later two at once, but they soon stopped. 

Feb. 23. Tonight at 8:50 P.M. when air was ;6 F., two ornate chorus frogs 
were calling at the east end of the "ornate area," and one above it at west end. 
Could not find them. Water was 5 or 10 warmer. 

Feb. 24. When air was 64, 8:50 P.M., about three P. strec\eri were calling. 
They generally are almost wholly out of the water. The one I watched was 
wholly so. Even when not croaking, the throat is overmuch swollen. This one 
was quite brown, with hardly a dorsal spot showing. 

March 24, Beeville, Tex. Air is resounding with Pseudacris n. clartyi. When 
we came to little creek, we heard peculiar note, went up one road and started 



TREE FROGS: HYLIDAE 

across soft field, but gave it up, followed up another road, and found the one 
peculiar beast to be a male Pseudacris strecl^eri. It was in the middle of the road. 
We drove on. Coming back, we heard another in the same place and later a 
lone voice near our camp. Is it a land call ? 

Feb. 9, 1932. We received 13 live Pseudacris strec^en males from our friend 
Albert J. Kirn, of Somerset, Tex. In a subsequent letter (Feb. 15) he spoke of 
them as follows: "I sent by express thirteen of the Pseudacris (ornata) street^- 
eri: two of them reddish (i cinnamon and smooth, the other warty) two 
green and the remainder normal colored. So you will find that at least color 
and smoothness of skin have nothing to do with the stability of the subspecies. 
The frogs quit calling the last night I got them, or at least I have heard very 
few since then. They can be heard clearly for a distance of more than half mile. 

"I had on hand three specimens of Pseudacris in pickle (one small). I in- 
cluded them with ten Scaphiopus hurteni that I mailed a few days ago." 

Notes on these three are: (i) A female 35 mm. long, practically devoid of 
dorsal and lateral spots or leg bars and with no vitta; taken at Lytle, Tex., 
Nov. 50, 1931, by A. J. Kirn. (2) A well-spotted spent female, 34 mm. long. 
Mr. Kirn's notes are: "Lytle, Texas, May 31, 1931. Dug into sandy soil about 
i% inches to its back. Found by our seeing the sand pushed up. Silent since 
January. Call all winter." Mr. Kirn unhesitatingly identifies these two as 
Pseudacris (ornata) strecfen. (3) A small, much-spotted, slim-legged speci- 
men 26.5 mm. in length, taken 1931 in Pleasanton, Tex., by L. Clayton Foster 
for A. J. Kirn. The latter merely identifies this specimen as Pseudacris. Were 
it not for the numerous dorsal spots, we might put it with the Pseudacris ornata 
of Florida, whose fingers arc less chubby than those of the adult P. strecl(en 9 
but this character, like the slender, prominently barred hind legs, may be an 
indication of its immaturity. We provisionally call it P. strecfyn. 

At last, March 24, 1932, we have in hand at the same time live frogs of the 
so-called form Pseudacris occidentals from Florida, and live frogs of the so- 
called form Pseudacris ornata from Texas. They are different forms. There is 
no doubt that the form from Florida is the Pseudacris ornata of Holbrook and 
the form from Texas is no longer P. ornata. We call it Pseudacris ptrecl(cri. 
Several live frogs of each form put side by side reveal the following from super- 
ficial examination : 

Pseudacris ornata from Florida is the more slender form: the snout is 
pointed; the arms and legs are longer and more slender; the fingers and toes 
are long and slender, with a mere trace of web at base of toes. The indistinct 
darker marks on the back are bars. The nostril is equidistant between snout 
and eye; the dark bar on the arm is long. 

Pseudacris strecl(eri from Texas is a short, fat, squatty form: the snout is 
shorter and broader at the tip; the arms and legs are short and broad; the fingers 
are short and fat, giving the frog a toadlike hand. The dark vitta on the face 
usually ends in front of the arm insertion. The dark rim of the upper jaw is con- 



280 HANDBOOK OF FROGS AND TOADS 

spicuous. The light area is broad at the nostril and again back of the eye. The 
dark pattern on the back is made of irregular, spotlike bars, these frequently 
having a fork at the upper end. The marks on the legs are only partial bars; the 
spots are inclined to be lighter in the center and darker at the rim even when 
indistinctly outlined with light. The bar between the eyes may be a conspicu- 
ous V. The dark spot is at the base of the arm only, or slightly on the upper arm. 

Authorities 9 corner: 
J. K. Strecker, Jr., Tex., 1915, p. 48. 
G. A. Moore and C. C. Rigney, Okla., 1942, p. 78. 

"Dec. 4th. A few Chorus frogs heard late P.M. yesterday and today late P.M. 
Heard late last night also. A few heard Nov. 18. Rainy at these times. 

"Dec. 6th. Chorus frogs calling last night when I went to bed. Temperature 
43, calling again tonight TO P.M. 45. 

"12/26. Chorus frogs calling every night. Heard at 10 A.M. and 5 P.M. yester- 
day. Weather cloudy, misty. 

"Jan. 19. Chorus frogs heard calling at noon, and all morning not so many 
but heard. Weather clear and mild. Rain day before and Thursday night. 

"1/27. Chorus frogs, a few calling now and then all day today. Weather 
cloudy, mild. 

"2/2. Many chorus frogs calling this evening and night. Weather milder. 
Temperature in evening, 48-53, misty weather for 3 days. 

"2/12. Chorus frogs also calling and today and Feb. 6, also Feb. 2, heard 
striped treefrogs (P. n. clarfoi) calling from water pools in south part of San 
Antonio. 

"2/19. Foggy all day; milder after 9 P.M. yesterday. Chorus frogs a few 
tonight. 

"2/24. Weather in afternoon a little cooler. Chorus frogs tonight and last 
night, not many" (notes, A. J. Kirn, Somerset, Tex., 1944). 

We named this species after Dr. John K. Strecker, Jr., for many reasons: 

(1) Dr. Strecker was a pioneer herpetological student in Texas life histories; 

(2) he did many personal favors for us and for other workers; and (3) he first 
described this form in his paper "Studies in North American Batrachology : 
Notes on the Robber Frog (Lithodytes latrans Cope)" (Trans. Acad. Set. 
St. Louis, June 14, 1910, XIX 5 , 73-82). 

[In 1910 few knew Pseudacris forms, and Strecker's statement that Litho- 
dytes latrans laid egg masses in water and had tadpoles in water seems, in 
season and description, to apply to P. strec\en. Doubtless Dr. Julius Hurter, his 
early herpetological mentor, helped him identify the frog, for the paper ap- 
peared in the St. Louis Academy of Science Transactions. He (Hurter) too 
must have been confused because in the USNM Hurter collection (USNM no. 
57706) we found P. strecl^eri labeled Lithodytes latrans. We have examined all 
the Baylor University material, and by 1915, certainly by 1926, Strecker knew 
the Pseudacris well and the fog over names was clearing.] 



Genus HYLA Laurent! 

Maps 18-24 

Anderson Tree Frog, Anderson Frog, Anderson's Hyla, Anderson Tree Toad, 

Green and Yellow Tree Toad 

Hyla andersonii Baird. Plate LIX; Map 19. 

Range: Central New Jersey (Middlesex Co.) to South Carolina. 

"About eight years ago James Chapin and I found this beautiful Tree Frog 
at the Runyon Pond two miles south of Sayreville, Middlesex Co. [NJ.|. Since 
that time I have found it at several other localities in the same region one mile 
south of Old Bridge, about one mile southeast of Browntown and at Freneau 
near Matawan. These localities are all in the sandy pine barren 'island* north 
of the Pine Barrens proper. Thus this species extends northward to within three 
miles or less of the lower Raritan River, its range coinciding at this point with 
that of the Carolina Chickadee" (W. DeW. Miller, N.J., 1916, p. 68). 

"Considerable doubt has been cast on the type locality record by later work- 
ers. Thus, the Anderson record 'has always bothered' Wright (1932), who re- 
marks: 'Anderson is twice as far from the coast as Southern Pines, N.C. (where 
he also took specimens) or Cheraw, S.C. is from the coast. Possibly the species 
extends that far inland along the Savannah River. . . . Anderson's tree frog 
is a Lower Austral (Austroriparian, Upper Coastal, etc.) form and Anderson, 
S.C. is in Upper Astral (Carolinian, Piedmont, etc.) country.' 

"Pickens (letter, 1938) points out that at the time the type specimen was col- 
lected most of the travel to the 'Up Country' was inland from the coast. Miss 
Charlotte Paine (later Mrs. M. E. Daniels), originally from Maine and an in- 
veterate collector, sent many specimens of amphibia, etc., to the Smithsonian 
Institution and she has generally been credited with taking the type of under- 
sonii. If this was the case, Pickens goes on to suggest, she may have picked up 
the specimen in question at some junction point, as Charleston, Columbia, or 
Augusta. 

"Although I believe that andersonii may occur in the Savannah River swamp 
well above the Fall line, the question can be satisfactorily settled only by the 
taking of another specimen reasonably close to Anderson" (E. B. Chamberlain, 
S.C., 1939, p. 21). 

Habitat: White cedar swamps. We found larvae in several pools, grassy, 
sedgy, sphagnaceous, along a dense woody border below one of the lakes at 
Lakehurst, N.J. 



282 



HANDBOOK OF FROGS AND TOADS 



"The exact positions of ten individuals were located, of which seven were 
captured. High-bush Blueberry tangles festooned with Green Briars made fur- 
ther investigations in this line impossible. The individuals are here referred to 
by number. 




Map 79 

"Numbers i, 2, 3, 4 and 5 were in one group, within 50 feet of the main 'pike' 
from May's Landing to Hammonton. The ground was covered quite evenly 
with Blueberry bushes from a foot to 18 inches high. Scattered Pitch Pines up 
to 12 inches in diameter stood from 10 to 30 feet apart. The ground was at 
most damp, and the only water nearby was a shallow pool about 30 feet away 
which probably dries up in the summer. Near the bases of the pines stood taller 
Blueberry bushes, up to three feet in height. No. i was sitting on the main stem 



TREE FROGS: HYLIDAE 283 

of a small Blueberry bush, 18 inches from the ground and six inches from the 
tip of the bush. A Pine stood i% feet away. No. 2 was on the leaf of a blueberry 
bush, 2% feet from the ground and 1% feet from a Pine trunk. No. 3 was on a 
little twiglet growing out from the trunk of a pine 3% feet from the ground. 
No. 4 was on the ground at the base of a Pine. No. 5 was one foot from the 
ground, where the twig of a Blueberry bush lay against the trunk of a Pine. 
All of the specimens in this group showed a strong preference for the near 
vicinity of a Pine. 

"Nos. 6, 7, and 8 were in a thicket of small Red Maples and high Blueberry 
bushes in a creek 'bottom/ No. 6 was on the main stem of a Blueberry bush 
about four feet from the ground. No. 7 was similarly located. No. 8 was about 
six feet from the ground in a small Red Maple. 

"Nos. 9 and 10 were in a thick tangle of high Blueberry bushes and Smilax. 
Both were near the tops of Blueberry bushes at least nine feet from the ground. 
For fifty feet around none of the vegetation was any lower, so it seems that these 
individuals climbed higher than is usual for the species in order to be out in 
the open. 

"Not all of the individuals were as tame as is generally noted for andersonii. 
A number of individuals would not continue singing when the observer turned 
the light on them or approached nearer than fifteen or twenty feet, and so could 
not be located. A silent andersonii in a thick tangle of Blueberry bushes could 
give points on hiding to a very small needle in a very large haystack. No fe- 
males were taken" (A. B. Klots, N.J., 1930, pp. 108-111). 

Size: Adults, 1^-1% inches. Males, 30-41 mm. Females, 38-47 mm. 

General appearance: This is a small green tree frog. The light-bordered, 
plum-colored band along the side of the body, with its yellow spots below, 
gives this beautiful little frog its distinctive character. This band marks its 
green dorsal color very sharply from its white ventral parts. It has orange in 
axilla and groin and on the rear of the femur. The throat of the female has a 
white-bordered green patch on either side. In its stout body it differs from the 
more slender and larger Hyla cinerea. 

Color: Male. Upper parts (dorsum, upper lip, angle of mouth, and dorsal 
surfaces of limbs) cress green to a light cress green, on sides lighter to deep 
chrysolite green. Stripe along side, behind vent, along limbs and upper jaw 
cartridge buff, ivory yellow, marguerite yellow, or seafoam yellow. Area back 
of eyes, along side to vent vinaceous-drab or purple-drab, becoming over tym- 
panum dark purplish drab. A little of the same color scattered on throat. Tops 
of forefeet and hind feet deep brownish drab except for first two digits, which 
have the cadmium yellow spots on the top. Fore part of underside of antebra- 
chium and under part of brachium with cadmium yellow, deep chrome, or 
orange spots; the same color also in axilla, groin and most of fore and hind 
parts of femur, whole undersides of tibia, and inner side of foot. All these 
orange or deep chrome spots are on a raw sienna or mars yellow background. 



HANDBOOK OF FROGS AND TOADS 




Plate LIX. Hyla andersonh (X rl /0- i~3 
Males. 



Iris more or less vinaceous-drab or 
purple-drab with vinaceous-tawny 
spots. Under parts white except for 
dark purplish drab of throat. 

Female. Throat white, grayish, or 
with slight purplish drab tinge. Green 
patch below angle of mouth usually 
white-edged, this usually absent in 
males. 

Structure: Form stout, head broad 
and flat; skin smooth; posterior sur- 
face of femur spotted ; vocal sacs sub- 
gular, 

Voice: This frog calls aquac\- 
aquack^aquacl{ t many times, perhaps 
twenty, at infrequent and irregular 
intervals. 

Dr. C. C. Abbott (N J., 1890, p. 189) 
says, "The andersonii utters a single 
note, better described by the syllable 
l keck/ which it usually repeats three 
or four times. It is not a frog-like note 
at all, but much resembles the call of 
the Virginia rail (Rallusvirginianus) ." 

"June 16-28, 1928. To the writer the 
call seemed a nasal 'quack/ almost 
verging on a 'quank' but without the 
strong V sound of the latter. The call 
was never disyllabic. 

"The note is repeated at about half- 
second intervals for sometimes fully 
30 seconds. When the frogs are in full 
song an interval of about two minutes 
intervenes between outbursts. We had 
no difficulty in starting the frogs call- 
ing again at distances of from fifty to 
three feet, after they had been silent 
for a minute or so. One individual 



was recorded as having called 74 times in one period of song. 

"The frogs definitely associate together for singing, whether because of the 
presence of females or for companionship. The latter probably plays a consid- 
erable share in the performance, as is evidenced by the quick response to an 
imitation of the call. Five such singing groups were definitely located. Of these 



TREE FROGS: HYLIDAE 285 

the first contained seven individuals, the second contained three, the third con- 
tained eight, the fourth contained three and the fifth, which was just across an 
uncrossable creek, contained at least six. Only once was a single individual 
noted in song alone, and that was a frog which called three times in a spot a 
half-mile distant from any others and was never heard from again. 

"The locations of the groups were fixed, and during our stay did not change 
a particle. Night after night a group would be in exactly the same area, though 
the individuals composing it shifted position a bit. 

"The time of singing was remarkably constant. On every night but one 
[June 16-28, 1928] the chorus started between ten and fifteen minutes before 
sundown. On the one exception, a clear dry night with a bright moon, the first 
songs were not heard until twenty minutes after sundown. 

"The carrying power of the song was excellent. A chorus was plainly heard 
as an entity over 800 paces away, with two patches of woods and a brushy 
swamp intervening. The wind was negligible. Individual voices were distin- 
guishable 754 paces away down a straight road, with a light wind blowing 
from the observers toward the frogs" (A. B. Klots, N.J., 1930, pp. 108-111). 

"At dusk we have usually taken our supper, and then waited for darkness 
to come on and for the Hylas to begin to sing; we have had good luck taking 
Hylas by the following method: One of us with an electric flashlight would 
start for the nearest singing Hyla, while the other waited some distance away. 
As soon as the Hyla stopped singing, the person who was not trying to ap- 
proach would imitate the call of the frog, and this would start it singing again 
vigorously, and while it was singing the collector bearing the light would ap- 
proach as quickly as possible, standing still as soon as the singing ceased. This 
process was kept up until finally the light flashed on the vibrating white throat 
of the singing Hyla, and its capture then became a perfectly simple matter, as 
they stared stupidly at the brilliant light. . . . This year, however, we did not 
get down to Lakehurst until the 8th of July, when we found the Hylas singing 
in goodly numbers in the white cedars about the lake. After capturing a num- 
ber of singing males (I had never taken a female before), my light flashed by 
merest chance upon a pair of Hylas sitting well up in a pine tree, in embrace. 
This, and another taken in a similar situation, were the only females secured, 
although we took several males from the low oak shrub about a small fresh 
water pool in the pine barrens" (T. Barbour, N.J., 1916, pp. 6-7). 

Breeding: This species breeds from May i to July 20. The eggs are single, 
attached to sphagnum or on the bottom. The egg is /4o-M inch (1.2-1.4 
mm.), the inner envelope Ki> inch (1.9-2.0 mm.), the outer H-% inch (3.5- 
4.0 mm.). The olive tadpole is small, i% inches (35 mm.), its tail medium 
long with tip acuminate. The tooth ridges are %. After 50 to 75 days, the tad- 
poles transform from the end of June to September i at %-% inch (11-15 
mm.). 

Noble and Noble (N.J., 1923) say that the eggs are single, scattered among 



286 HANDBOOK OF FROGS AND TOADS 

waterweeds, attached to sphagnum, or free on the bottom, in small, nonstag- 
nant pools or in slow-moving streams of the pine barrens. Color, dark brown 
and creamy white. Egg 1.2-1.4 mm - Inner envelope 1.9-2 mm. Outer envelope 
3.5-4.0 mm. Complement 800-1000. 

"When the tail is nearly absorbed and they leave the water, they are about 
25 mm. long and of a dull olive green. They grow lighter, that is, brighter green 
in hue with the disappearance of the tail, until the little frogs, which in length 
of body are 15 mm., resemble the mature individuals. The white that margins 
the green of the back and extremities is not so conspicuous as in the adults, and 
the saffron of the underparts is wanting in those that I have examined" ( W. T. 
Davis, N.J., 1907, p. 50). 

Journal notes: On June 8, 1922, on our way to Okefinokee Swamp, we 
camped in the evening on the north side of Everett's Pond, N.C., near the 
S. Car.-N. Car. line. The instant it became dark we heard plenty of Acns gryl- 
lus, Bufo jowleri, and Rana catesbeiana, a few Rana clamitans and Hyla cinerea 
in chorus, and Hyla versicolor. . . . Later heard Hyla andcrsonii. These H. 
andcrsomi were in a stream or branch with sweet gum, tangle of oaks, bamboo 
briars (Smilax), Magnolia glauca, maples, black gum. Later in the evening 
Miles Pirnie heard one opposite our camp. All the frogs heard were in or near 
the lake. Harper speaks of the calling of these Hylas thus: "I heard the note 
of H. andersonn at a distance of 200 yards and suspected almost at once what 
it was. Trailed it and after a long wait located it in some tall bushes in the edge 
of a branch swamp. Its note bears a general resemblance to that of H. cinerea 
but goes about twice as rapidly, is about half as loud and sounds more like quak, 
quak than quonk. It carries tairly well at 200 yards and about 300 yards would 
probably be the limit. . . . Did not see it croak. Its periods are infrequent and 
irregular, 2 minutes' interval. Perhaps 15 or 20 or 25 notes given in one period." 

On June 28, 1929, we heard these Hyla andersonn in the wooded edge of 
Mr. P. H. Emilie's lake at Lakehurst, NJ. The males were among magnolias, 
maples, huckleberries, azaleas, Ilex glabra t Viburnum. 

Authorities' corner: 

W. T. Davis, N.J., 1905, p. 795. C. S. Brimley, N.C., 1940, p. 22. 

W. T. Davis, N.J., 1907, p. 49. L. R, Aronson, N.J., 1943, pp. 246-249. 

G. K. and R. C. Noble, N.J., 1923, pp. 

419 ff. 

H. andersonii is a handsome species and it has been superbly illustrated. 
From the time of Miss Dickerson's beautifully colored plate in her fine Frog 
Boot^ (Chapin's specimens 1906-1908), and L. A. Fuertes' painting of them to 
the days of our private preparatory tips to Dr. Noble's well-illustrated, first 
(1923) outdoor life history of H. andersonii, we have been interested in this 
form. Several students, Mr. and Mrs. A. B. Klots, W. H. Caulwell, and others, 
have brought us specimens, particularly in the period since 1923. 



TREE FROGS: HYLIDAE 287 

Canyon Tree Frog, Canyon Tree Toad, Desert Tree Toad, Cope's Hyla, 
Arizona Tree Frog, Sonoran Tree Toad 

Hyla arenicolor Cope. Plates LX, LXI; Map 20. 

Range: Southwestern United States : western Texas through extreme south- 
ern Colorado and Utah to the region of the Colorado River in Nevada, down it 
to Lower California and up southern California!! coast to Ventura County. In 
Mexico to Guadajuata, Guadalajaca, and Toluca. 

Habitat: Rocky canyons. From Fern Canyon in Ft. Davis Mts., Tex., to 
Tahquitz Canyon, Calif. Invariably we found them in rocky, clear streams. 
They have been reported by others from mountain springs, irrigation ditches, 
and "rapidly flowing streams." 

"Throughout this region [Southern California], its habitat appears to be 
confined to streams and mountain springs between 1000 and 5000 feet eleva- 
tion. Here the writer has found it associated with such trees as Alnus rhombi- 
jolta, Platanus racemosa, and Acer macrophyllum and in this state at least it 
may be regarded as an inhabitant of canyons within the upper sonoran zone. 

"It is apparently more strictly aquatic than the smaller Hyla regilla Baird 
and Girard, whose range in Southern California is, in part, co-extensive with it. 
The former species has never been found far away from the vicinity of water, 
while the latter has often been seen under vegetation a considerable distance 
from it" (C. H. Richardson, Jr., Gen., 1912, p. 607). 

"Occupies chiefly the Upper Sonoran life-zone, extending locally into Lower 
Sonoran. Lives on boulders and exposed rock faces close to canon streams" 
(J. Grinnell and C. L. Camp, Calif., 1917, p. 145). 

"Fairly common around shady moist places in the Canyon and occasionally 
up to South Rim. Lower Sonoran and Upper Sonoran Zones" (E. D. McKee 
and C. M. Bogert, Ariz., 1934, p. 178). 

"Common in the pine-fir and chaparral-woodlands zones but uncommon in 
the semidcsert. Calling individuals and clasping pairs were common in water 
at a dam in the pine-fir forest on May 19, 1936. After a mating season adults are 
found occasionally about buildings" (E. S. Little, Jr., Ariz., 1940, p. 262). 

"This species is confined to the High Plateaus-Colorado River region of 
Utah. They are common in Zion National Park and St. George. . . . This 
species is commonly found on the rocks and banks of the small streams and 
pools of the Colorado River drainage of this state. It is seldom if ever found 
farther than a few feet removed from running water" (V. M. Tanner, Utah, 
1931, pp. 186-187). 

Size: Adults, i%-2% inches. Males, 29-5? mm. Females, 30-54 mm. 

General appearance: This tree toad averages smaller in size and duller in 
color than our common tree toad (Hyla versicolor). When sitting quietly it 
(a specimen from Grand Canyon, Ariz.) looks surprisingly like the little 
spotted canyon toad (Bufo punctatus), but when it climbs, its conspicuously 



2 88 HANDBOOK OF FROGS AND TOADS 

long legs and large disks on fingers and toes immediately show it to be a tree 
toad. It easily clings to a vertical surface and climbs up even a sheet of glass. 
The back is brown or grayish in color with dark dots scattered over it. The legs 
are barred with dark areas. There is much yellow or orange on the rear of the 
legs, in the groin, and in the axilla. It is light beneath. The skin is slightly 
rough, becoming more so as it becomes dry. Unlike the Pacific tree frog (Hyla 
regilla), this species has no stripe through the eye and along the side of the 
body. The skin is roughened in H. aremcolor, smooth in H. regilla. This frog 
has slight webs between the fingers. 

"Size small, head-and-body length 50 mm. (2 inches) or less; fingers and 
toes with expanded adhesive disks; no webs between fingers; dorsal skin 
rough-surfaced, with many small papillae; side of head concolor with rest of 
head and body; coloration light or dark grny. . . . Distinguished from other 
Salientia of California by expanded disks on fingers and toes and by small size 
of adults; from Hyla regilla by average larger size (especially of females), by 
larger discs on digits, by rougher skin on dorsal surface of body, and by lack 
of dark stripe on side of head through eye" (T. I. Storer, Calif., 1925, p. 204). 

Color: Scntenac Canyon, Calif., from L. M. Klauber, March 31, 1928. Male. 
Upper parts olive-huff, drab, light grayish olive or deep olive-buff. Snout ahead 
of eyes avcllancous or wood brown, some of larger warty spots with touch of 
same color or pale cinnamon-pink to vinaceous-cinnamon. Bar between eyes, 
line in front of eye and from eye through ear, four or five larger spots on back, 
three crossbars on femur, four on tibia, four on hind foot, and two on forearm 
are from deep olive-buff to citrine-drab. These spots are outlined with broken 
black and filled in with body color giving the deep olive-buff or citrine-drab 
combination. Interspaces between bars on legs and arms vinaceous-buff or avel- 
laneous. Groin, olive-ocher, yellow ochcr, lemon chrome, becoming aniline 
yellow or raw sienna on front and rear of femur. Dorsal color on femur very 
narrow. Dorsal color extends around vent; with tilleul buff or pale olive-buff 
on each tubercle. Ventral view of legs and arms vinaceous-fawn or light russet- 
vinaccous or light grayish vmaceous. Venter white or cartridge buff. Vocal sac 
when inflated light buff except near chin where spot below eye is light pinkish 
cinnamon or sulphine yellow. Iris with benzo brown, cinnamon-drab, or 
brownish drab band in front and back of pupil, this finely dotted with olive- 
buff; iris below and above pupil largely olive-buff with wavy, more or less 
vertical lines of brownish drab; pupil rim above dull green-yellow or chalced- 
ony yellow, light below; pupil rim below with vertical slit of black. Another 
male has back pale olive-buff or pale drab gray. 

Another male. Dorsal color pale olive-gray or pale smoke gray to light 
grayish olive with light vinaceous-cinnamon on snout and forelegs. Belly not 
white but dark olive-buff. 

Female. Underparts white or pale olive-buff, middle of belly or rear belly 
olive-buff. Groin with very little lemon chrome. 



TREE FROGS: HYLIDAE 



289 





Plate LX. Hyla arenicolor. (XO- Part 
i. 1-5. From Grand Canyon, Ariz. 6,7. 
Transformed and tadpole, from Fern Can- 
yon, Davis Mts., Tex. 



Plate LXL Hyla arcnicolor. (XO- Part 
2. From California. 



2 9 o HANDBOOK OF FROGS AND TOADS 

Another female. Fern Canyon, Tex., July 9, 1925. Spots on back, body, 
legs, head mouse gray to dark olive-gray; back deep grayish or olive becoming 
pale smoke gray on the sides next to belly color, on labial margin, snout, and 
.vent region. In axilla, side of lower belly, front and rear of thigh, rear of tibia 
and inner face of foot, rear of humerus, somewhat on inner face of forearm 
olive-ocher or aniline yellow or sulphme yellow, becoming on the edges green- 
yellow. Breast white. Throat white spotted with mouse gray or dark olive gray, 
especially around edge of lower jaw. Tympanum with some orange rufous or 
tawny in lower half. 

"The color phases run from light gray to brownish black, with numerous 
roundish spots irregularly distributed and not forming well-marked patterns, 
except on the hind legs, which appear banded. In the light and dark color 
phases the spots become obscure" (G. P. Engelhardt, Utah, 1918, p. 79). 

Structure: Long arms and legs with large disks on tips of fingers and toes; 
slight webs between the fingers; skin somewhat roughened; more or less uni- 
form in color; tympanum small; prominent fold across breast; prominent fold 
on tarsus. 

A. A. W. has long held it likely that there is more than one race of Hyla 
arenicolor. The following observations are some of her notations: In general, 
the frogs from California have longer legs and a black bar across the head, a 
lighter gray background with inky spots, and a more pebbly skin. The frogs 
from Grand Canyon, Ariz., are smoother, have shorter legs, arc reddish in 
color, spotted rather indistinctly, and bear no marked triangle on the head. 
Those from Rainbow Bridge collected by Eaton are like the Canyon ones. 
Those from the White Mountains are rather pebbly, and the one from New 
Mexico is pebbly. The Canyon frogs had larger disks on the toes, larger eyes, 
and less distinct spots. In life they sat with body parallel to finger when held, 
in contrast with the more erect, toadlike posture of the California frogs. 

"Hyla affinis, Baird. Body rough. Tympanum two-thirds the size of eye. 
Tibia not quite half the length of the body, but reaching more than halfway 
from anus to center of eyes. Color ash gray or green, with numerous rounded 
dorsal blotches. Three transverse bands on each thigh and leg. No vermicula- 
tion on anterior and posterior faces of hind legs, nor on lower part of sides. A 
light spot under the eye. Web of hand extending only to the third joint of the 
second finger. Arm from elbow less than tibia, but longer than hind foot. 
About i% inches long. Hab. Northern Sonora" (S. F. Baird, Gen., 1854, p. 61). 

"General aspect of H. versicolor, having the same squat appearance, the 
granulated skin above and below, the ash-color back with darker mottlings, 
the white spot under the eye, etc. The most conspicuous distinctive features are 
the absence of webs ot the fingers, the greater length of the hind legs, and the 
blotches on the back being in round spots, not cuneiform. The legs with three 
bars not two, and without the reticulate markings behind and below" (E. D. 
Cope, Gen., 1889, p. 369). 



TREE FROGS: HYLIDAE 291 

Voice: This call has been described as the "quack of a duck" by Storer, and 
as "the bleat of a goat" by Duges. To us the note sounded lower, much weaker, 
and less blatant than that of Hyla versicolor and not so persistent. "It was like 
a hoarse sheep's 'Ba-a-a' when they were in the open, but more like a roar when 
they were under the rocks" (T. H. Eaton, Ariz., 1935^ p. 7). 

"The frogs are most vociferous in the late afternoon and rarely are heard at 
night" (G. P. Engelhardt, Utah, 1918, p. 79). 

"The song of this tree toad is one of the most melodious of all our species" 
(V. M. Tanner, Utah, 1931, p. 187). 

"The note differs markedly from that of our common western treetoad 
(Hyla regilla) in being lower in pitch, somewhat weaker in volume, and with- 
out any tendency toward a two-syllable sound such as is heard often from 
regilla. On two occasions, in San Diego County, the two species were heard 
side by side and there was not the slightest difficulty in distinguishing them. 
Two separate individuals of arenicolor which were timed by the watch were 
croaking at one-second intervals. With all the males noted on Tahquitz Creek 
(fully 25 were located) only one female was seen. The chorus began about 
4 P.M., soon after the sun had disappeared behind the San Jacinto Range, was 
strongest just after dark, from about 7:30 to 8:30 P.M., and continued on into 
the night at least until 2 A.M. On other occasions males of Hyla arenicolor were 
heard croaking up to about 6 A.M., but only a few notes were given as late as 
that hour in the morning. The duration of the 'song' season is unknown; on 
the morning of June 21, 1919, Canon Tree-toads were heard croaking in canons 
near the Mount Wilson trail" (T. I. Storer, Calif., 1925, p. 208). 

"Tree toads of this species were found in White House Canyon, in the Santa 
Rita Mountains, August 3. Males were heard calling just before dark from 
branches of oak trees, about fifteen feet from the ground. The call was an even, 
strong trill" (F. W. King, Ariz., 1932, p. 176). 

Breeding: This species breeds from March i to July i. The eggs are single, 
floating near the surface or on the bottom of the pool usually attached to leaves 
(Atsatt and Storer). The tadpole is medium in size, 2 inches (50 mm.) long, 
dark olive in color with some tail crests suffused with reddish, orange-pink to 
coral red. The tooth ridges are %. After a period of 40 to 75 days, the tadpoles 
transform from June i to August 15, at % inch (14-17 mm.). We have seen 
mature tadpoles collected in April from Fern Canyon, Ft. Davis Mts., Tex., 
and neither in 1917 nor in 1925 did we hear the choruses after July i, and rarely 
single voices. C. H. Richardson, Jr. (Gen., 1912, p. 607), indicates "that the 
breeding season . . . extends from late spring until fall." The extension of 
the breeding season to fall is apparently based on one female taken Sept. 4, 
1906, and containing large eggs. 

"White House Canyon, in the Santa Rita Mountains. . . . Tadpoles of 
H. arenicolor were found in pools nearby with some of the larvae transform- 
ing at 18 mm. body length" (F. W. King, Ariz., 1932, p. 176). 



292 HANDBOOK OF FROGS AND TOADS 

"I have found it to be fairly common at the Water Cress Spring in St. George 
in February and on August 28, 1919 1 found newly hatched as well as matured 
tadpoles in the 'Stadium pool' in Zion National Park. The tadpoles are black 
when small becoming grayish when older" (V. M. Tanner, Utah, 1931, pp. 186- 
187). 

"Ova were observed in the form of small clusters deposited along the margin 
of pools. The tadpoles, at first black, later become mottled gray when they re- 
sort to deeper water" (G. P. Engelhardt, Utah, 1918, p. 79). 

Journal notes: July 16, 1917, Pinaleno Mts., Ariz. On the rocks over the wa- 
ter, Anna put her hand accidentally on a Hyla aremcolor and Ray Shannon 
found another on a tree trunk about 4 feet up. Paul Needham found another 
on a boulder just above the water. They do not seem at home in the swifter 
water. 

July 5, 1925, Fern Canyon (near Alpine, Tex.). In the ravine just after a 
drenching ram, we found four transformed Hyla aremcolor on the boulders 
some 25 or more feet above the level of the creek. The creek was full of them 
before the rain. Today, July 6, the creek is down and clear. Near the falls we 
often found one or two Hyla aremcolor tadpoles amongst the boulders in shal- 
low water or swimming at the surface. We secured today only one mature 
tadpole. The rest were small tadpoles. At the pool below the falls were many 
of them. . . . Also some above the falls. 

July 7, Ranger Canyon. About 6:45-7 P.M., we arrived at the big pool or 
falls of cast Ranger Canyon. Mr. L. T. Murray espied a frog in amongst the 
rocks in our very midst. It was an adult female Hyla arenicolor. 

July i i, Toyahvalc. We drove to Madera Canyon for a night camp. ... In 
the canyon caught a recently transformed Hyla aremcolor, and just about dark 
heard several males calling. 

June iH-2i, 11^4, Pcna Blanca Spring, Ariz. Found two Hyla aremcolor in 
the overflow area around the spring, 

April 26, 1942. In Santa Anita Canyon (see R. b. muscosa), Calif., beside the 
river we caught two Hyla aremcolor. The first one I spied was on gravel be- 
side stream and hopped in. It was very light in color much granitic rock here, 
light gray in color. Almost no sediment in stream. 

April $o. At mouth Strawberry Creek, Calif., above bridge there leaped in 
from a boulder a female H. aremcolor. 

May 2. North of Rmcon at about 2000 feet in a little stream beside the road, 
shaded by live oaks, found perched on a boulder, 3 inches from each other, a 
male and female H. aremcolor. They were in full sunlight. 

May 6, Buckman Spring, Calif., to bridge 3*4-4 miles south. During the 
evening rode north toward Buckman Springs, hearing several isolated calls 
which took me some time to locate. Usually they would quit when I ap- 
proached. One was in a bush. Finally on a boulder upside of stream current 
saw one with bubble throat inflated. Hyla arenicolor. Must have been ten 



294 



HANDBOOK OF FROGS AND TOADS 



males scattered along the creek. Their call is not long, not so grating as that 
of H. regilla, neither loud nor strident. No such voluminous call as Hyla 
versicolor. 

Authorities 9 corner: 

G. P. Englehardt, Ariz., 1917, pp. 5-6. J. R. Slevin, B.C., 1928, p. 112. 
S. R. Atsatt in T. I. Storer, Calif., 1925, L. T. Murray, Tex., 1939, p. 5. 

p. 209. 

Viosca's Tree Frog, Whistling Tree Frog, Whistling Frog, Bird-voiced Hyla, 

Bird-voiced Tree Frog 

Hyla avivoca Viosca. Plate LXII; Map 21. 

Range: Southeast Louisiana (Florida parishes of Louisiana) to Florida 
(O. C. Van Hyning) ; Georgia (F. Harper) ; Henderson, Tenn. (Endsley) ; 
Kentucky (Parker). 

Habitat: Tupelo swamps in the valleys of rivers and smaller streams on tu- 
pe^o or cypress trees and on buttonbush (adapted from Viosca, 1928). 

"The Apalachicola drainage. Known from Jackson and Liberty counties. 
Habitat Tupelo, titi and cypress swamps. Not rare, but infrequently seen be- 
cause of the nature of its habitat" (A. F. Carr, Jr., Fla., 1940^ p. 58) . 

Size: Adults, i%-i % inches. Males, 28-39 mm. Females, 32-49 mm. 

General appearance: These frogs are a small edition of our common tree 
toad, Hyla v. versicolor. They may be brown, green, or gray, light or dark. 
The arms and legs are distinctly barred. There is the characteristic light yellow 
or white spot below the eye. The color in the groin and on the rear of the legs 
is a pale yellowish green, instead of orange as in Hyla v. versicolor males. The 
throats of these males are more or less darkened with black specks. The dark 
pattern on the back consists of a bar across the head and the eyelids and an 
irregular area on the back, the bulk of the pattern being near the rump. The 
skin is moderately smooth, in some being very finely granular. In appearance 
they are more slender than H. versicolor, and when dark brown, as we first 
saw them, they reminded us strongly of Hyla fetnoralis. 

Color: Male. Top of head rainette green, courge green, or Scheele's green, 
with same color on upper eyelid and back to mid-back spot, and on either side 
of this spot. Interspaces of hind legs night green, courge green, or rainette 
green. The mid-dorsal spot appears dark olive or deep olive. Under the lens its 
background is black, spotted with apple green. An oblique bar of similar color 
extends from upper eyelid toward meson but does not meet its fellow. The 
color just above the vent may be clear yellow-green. One band across middle 
of femur is chamois or vinaceous-buff. A line of mummy brown separates dor- 
sal color of hind leg from ventral color. Outer half of rear of femur and rear 
half of dorsum is spotted with light lumiere green. Bars on tibia green like 
back of body. Bars on tarsus and foot with cream-buff. Spot below eye is pale 



TREE FROGS: HYLIDAE 



295 



glass green. Back of eye is a broad black vitta extending to shoulder. It sur- 
rounds the tympanum. From this vitta to groin is a row of black spots sepa- 
rating dorsum from venter. Tympanum is more or less the same color as femur 
bar or interspaces on top of arms and hands, which are almost old gold. Sub- 
orbital spot is margined with mummy brown or black. The forward part of 
chin is marked with black dots below the lower labial border. Forward throat 



+Hyla avivoca 9 1 




Map 21 

is white with a wash of marguerite yellow. Lower throat is pale flesh to Vene- 
tian pink. Lower belly and lower surface of femur are the same. Rear of femur 
below vent and rear of tarsus are marked with white on pebbly surface. Lower 
belly and groin have wash of viridme green or other yellow-green. Underside 
of fore limb is like fore part of venter, lichen green or light gull gray. Pupil rim 
of eye citron yellow or strontian yellow; ins black, heavily spotted with these 
yellows. 

Female, Mariana, Fla., from O. C. Van Hyning, Jan. 3, 1933. Top of head 
drab, buffy brown and the same color on forward part of dorsum. The bar 
from eye to eye, the forward dorsal spot, and the rear dorsal spot olive-brown. 
The side of body has background pale olive-buff to pale vinaceous-fawn, the 



296 HANDBOOK OF FROGS AND TOADS 

groin olive-buff or pale glass green. The light interspace color on rear of femur 
may be glass green, pale glass green, or pale olivine. The transverse bars on 
tibia, interspace color on rear of femur, and spots on groin are bister or raw 
umber. The pale spot below the eye is clear, prominent, pale olive-buff or 
marguerite yellow. The whole chin is white with very few black dots near 
lower labial border. The under parts in general are white or pale olive-buff, 
with scarcely any suggestion of the wash of greenish present in the male. The 
pupil rim is encircled by a carnelian red or vinaceous-tawny ring; the iris 
proper is black dotted with testaceous or terra cotta spots. 

Structure: More slender than Hyla versicolor; muzzle truncate; webs large, 
but leaving the last two joints of fourth toe free, except for web margin; back 
almost smooth; ventral surface with distinct granulations; usually with nar- 
row waist; male with folds on rear of throat almost to pectoral region. 

Voice: "The voice is birdlike, being a plaintive whistle repeated in quick 
succession, much as in the redbellied woodpecker. This call is sometimes pre- 
ceded by a few notes of a slower call much like the voice of Hyla crucifer. 

"The pitch and tone of the voice of avivoca is nearer to that of crucifer than 
to any other eastern American Hyla although its rate is far more rapid than 
that of crucifer" (P. Viosca, Jr., La., 1928, pp. 90-91). 

"The males call from bushes in or near the water and from cypress, tupelo, 
and black-gum boles" (A. F. Carr, Jr., Fla., 1940^ p. 58). 

See F. Harper, Gen., 1935, pp. 290-293, for characteristics of its call. 

Breeding: This species breeds from June to mid-August. The eggs and tad- 
poles are not described. "Hyla avivoca is more clearly related to Hyla versicolor 
than to any other North American Hyla. . . ." 

"I located a small chorus in Sweetwater Creek Swamp, Liberty County, 
April 19, 1935, where H. crucifer, H. v. versicolor, Pseudacns jenartim, and 
Rana clamitans were also calling. I have never seen a female" (A. F. Carr, Jr., 
Fla., i94ob,p. 58). 

Journal notes: June 8, 1930, New Orleans, Sunday. Went to Mandevillc and 
Lewisburg by myself to search out Viosca's type locality. Didn't succeed. 

June 9, New Orleans. Went to Viosca's house. Heard H. c. ctnerea. While 
there Chase from office called up that by dint of hard work he had taken four 
Hyla avivoca at Lewisburg, Covington, and in a swamp just south of Mande- 
ville. They are males. They surely are a different frog. 

June 10, New Orleans. Started at noon for the north country, Pearl River, 
and pine barren parishes of Louisiana, with Percy Viosca, Jr., and H. B. Chase. 
Went to Pearl River. Didn't hear H. avivoca in the gum and cypress edges, a 
haunt of the species. Presently we heard a Flycatcher, which I remarked was 
like a crested Flycatcher. Then Chase got excited over a note in the tupelo 
swamp either side of the bridge. Instantly the setting and note made me say, 
"It is a Pileated Woodpecker." A day or so before Viosca and Chase had said 



TREE FROGS: HYLIDAE 



297 




Plate LXII. Hyla avivoca (y(iVi). 1-4. 
Males. 



Plate LX1II. Hyla baudinii (X%). i. Fe- 
male. 2-7. Males. 



298 HANDBOOK OF FROGS AND TOADS 

this frog's note was like a Pileated Woodpecker at a distance. Naturally they 
both laughed at me. Worth it. Would I have guessed it to be a frog, much less 
a Hyla? Fear not. Heard a few of them in the gums of Tick Faw River where 
we will go tomorrow night. 

June ii. Tick Faw River, La. (near Ferry). In Tick Faw River we heard a 
few H. avivoca. Finally Chase called me to hurry and started off on the run. 
I followed. We were soon in a tupelo swamp. Near the edge we heard them 
Chase found his first one crosswise of leaves of buttonbush (Cefhalanthus 
occidentalis) . My first was on the bole of a gum. Many of Chase's captures 
were head down on upright branch of buttonbush. Mine were head up. Chase 
and I followed voices. We got five frogs between us. Viosca with a lamp on 
his head shone their eyes. He didn't follow voices so much. He got more in 
this way than either of us. One frog was pure green with no markings. Viosca 
found several on small gums near the tupelo edge while Chase's and my cap- 
tures were farther in the swamp. On the trees was some poison ivy. On clumps 
or tree bases in the swamp and at the edge of the swamp were sensitive fern 
(Onoclea senstbilis), royal fern (Qsmunda regalis), Utrictdaria (purple), a 
marsh St.-John's-wort, sphagnum, water penny (Hydrocotyle), lizard's-tail 
(Saururus), Nyssa aquatica, and buttonbushes with mayhaws. Most of the 
frogs were 3 or 4 feet above the water. Finally Chase found a female on a log, 
in the water. Sometimes they go higher in trees. Usually they are down at 3-4 
feet or less. Their call is birdlike. It is a fine, delicate little species. Viosca did 
right to call it a new form, H. avivoca. 

June ii. Tick Faw River, La. Throat in croaking is somewhat swollen, vi- 
brates for 6-10 or 12 calls. These are more or less the same, repeated. Sometimes 
raises rear end with calling. Chase also noticed the vibration or varied swelling 
at each call. 

Authorities 9 corner: 
P. Viosca, Jr., La., 1928, p. 91. 
J. R. Endslcy, Tenn., 1937, p. 70. 

Mexican Tree Frog, Van Vliet's Frog 

Hyla battdinii (Dumcril and Bibron) . Plate LXIII; Map 20. 

Range: South-central and southern Texas through Central America. 

"This Mexican species has been found by Mr. Marnock in the low country 
southwest of San Antonio, commencing with San Miguel Creek, a tributary 
of the Medina. This is its most eastern known range, that previously given by 
Professor Baird being Brownsville (as Hyla vanvlietif)" (E. D. Cope, Tex., 
i88oa, p. 29). 

Habitat: June 18, 1930. At 10 P.M., two miles west of Brownsville, in a resaca, 
found these frogs in small bushes, in weedy clumps, and even in grassy tangles 
in overflowed tomato field adjoining the overflowed resaca. Later, along the 



TREE FROGS: HYLIDAE 



299 



Rio Grande in Brownsville, found them in a pond. Mr. Blanchard tells me he 
saw Mr. Camp take them along the river (Rio Grande) in palms just above 
Mr. Rebb's palm grove. 

June 19. Heard several in the palm grove which is completely flooded by 
high river. Beyond grove, and along river, a large chorus is calling. 

Size: Adults, 1/4-3% inches. Males, 44-71 mm. Females, 44-89 mm. 

General appearance: This large tree frog has a black patch over the arm 
insertion and a white line encircling the arm insertion. Its color ranges from 
nearly black to light yellow green, gray, or fawn. It has a transverse bar be- 
tween the eyes. Various irregular lateral spots of black form a reticulation on 
a yellow or olivine side area. There are transverse bars on the legs, a light 
greenish spot under the eye, a light line above the dark-edged upper jaw, and 
a dark line from eye to shoulder ending in a prominent black patch. The 
breast and throat may be spotted with dark. The sides are yellow posteriorly. 
In the female, the under parts of the throat and upper breast are white, the 
lower breast and belly white suffused with pale green. In the male, under 
parts arc cream buff with lateral throat sacs, a pale brownish drab. The rear of 
the femur is yellow and russet. 

Color: Male. Brownsville, Tex., June 19, 1930. Back sulphine yellow and 
aniline yellow on side or olive lake on back becoming old gold on sides (no 
dorsal marks; last night this frog was brown with prominent marks on dor- 
sum). The back may be chamois to primrose yellow with olive-yellow or 
some green interspersed. Cream buff line along jaw to beyond angle of mouth 
and under tympanum. Tympanum pinkish cinnamon or light vinaccous- 
cinnamon or vinaceous-buff. From eye over tympanum irregular black line 
ending in a large vertical curl of black above arm insertion. Along side and 
a little below level of black arm insertion spot are several prominent black 
spots to the groin. This black spotting is more or less reticulated with lemon 
yellow. Top of fore limbs warm buff, pinkish buff, or chamois. From eye al- 
most to black shoulder spot is a wood brown band. Chin sulphur yellow or 
light green-yellow. Each lateral sac with oil yellow to ecru-drab or pale brown- 
ish drab. Under parts cream buff except a narrow pectoral area and extension 
between two sacs. This little area is white. Hind legs more or less color of 
forelegs. Rear of femur with some viridine yellow or bright green-yellow with 
vinaceous-russet; under side of hind legs onion-skin pink with every tubercle 
crowned with sulphur yellow. All of venter very pebbly. Iris very bronzy; 
below heavily spotted with buff-pink, above it is chartreuse yellow; pupil 
rim pale greenish yellow, broken in front and behind; outside iris is ring of 
lichen green or Niagara green. 

Other males. One greenish yellow on back except for top of head and top 
of hind legs and forelegs, which are viridine yellow. Another bright green- 
yellow on back. From eye of each extends a horizontal black line ending in a 
vertical black curve back of axil. Below this is a clay color or cinnamon band 



300 HANDBOOK OF FROGS AND TOADS 

outlined by black above, and below by light line from upper jaw edge almost 
to base of vertical black bar. Tympanum may be vinaceous-cinnamon or cin- 
namon. Along side where green of back ends and before greenish yellow of 
reticulated or spotted side begins is cinnamon-buff. In one, no black spots and 
little yellow. Iris, upper part cinnamon-buff. 

Female. Brownsville, Tex., from H. C. Blanchard, July 23, 1930. The for- 
ward part of head, upper eyelid, face, and arms are vinaceous-buff. The back- 
ground of back and legs is deep olive-buff or olive-buff. The bars of tibia, 
femur, and arm, the dark spots on back, and the large spot from each eyelid 
on fore back are grayish olive or vetiver green. Particularly in the groin and 
slightly on sides are black spots on pale olivine ground. The reticulation of 
the sides is less prominent than in the male, and restricted almost entirely to 
the groin. The black U-shaped spot on shoulder has the opening toward the 
tympanum. The tympanum is wood brown. The iris is somewhat duller than 
in the male; lower ins wood brown, the upper avellaneous. The under parts 
of hands are light vinaccous-fawn. The under parts of feet, femur, and tibia 
are light russet-vinaceous. The under parts of throat and upper breast are 
white; the lower breast and belly are white with a suffusion of pale turtle 
green. After the frog had been in a moist jar for a few hours, the back became 
darker, Prout's brown, and the reticulation along the sides became more 
marked. 

Aug. 13, the female still continues with the rich brown coloration. She re- 
mains under the wet moss. The two males come up in the glass jar, one is a 
bright green and the other old gold or citrine. Female 54 mm. 

Structure: Skin, smooth above, set with fine tubercles; under parts granu- 
lar; tympanum nearly as large as eye; fold of skin from eye to shoulder; 
prominent fold across breast; disks large, fingers slightly webbed, toes 
webbed; a distinct tarsal fold. 

"Hyla vanvlieth Baird Nearly smooth above. Tympanum nearly as large 
as the eye. Tibia half as long as the body, longer than arm from elbow, which 
in turn exceeds the foot. Ash gray or olive, with an irregular cruciform dorsal 
blotch. Thigh and leg with three transverse bands each. Their inner surfaces 
when flexed scarcely reticulated, but spotted with white upon a darker 
ground. Inside of tibia uncolored. Body two inches. Hab. Brownsville, Texas" 
(S. F. Baird, Gen., 1856, p. 61). 

Voice: The vocal sac is better developed on either side of throat than in its 
middle. This is especially noticeable in the live frog. 

June 18, 1930, Brownsville, Tex. The first time I heard them I guessed they 
must be H. baudinti. Heard their chorus at % mile. The note was a blurred 
Aec^ or l(ecl( (not high like Acris). This may be repeated 5 to 8 or even 10 or 
12 times. Then comes an interesting chuckle, or no chuckle at all, or no fecl^ 
and all chuckle. Never heard anything like it. The first call (not chuckle) in 
some ways reminds me of Hyla cinerea, but yet it is much unlike the call of 



TREE FROGS: HYLIDAE 301 

the cowbell tree frog. Does the full dilation of the sac on either side cause the 
queer note (unlike most Hylas) ? 

In Mr. F. Rebb's palm grove heard a few H. batidinu. They are too far out 
in an overflow to reach them. To the east of his house and near the river heard 
a chorus of these frogs. Their repeated fecfo or hect(s last about 2 or 3 seconds, 
then as much interval, whereas B. vdliceps has a longer call, sometimes to 5 
seconds. 

Breeding: The only record on transformation is of one specimen trans- 
formed, or just past, from Panama, % inch (21 mm.) in length, which was 
caught Feb. 19, 1911. They were in full chorus in Brownsville, Tex., June 19, 
1930. In southern Vcra Cruz, A. G. Ruthven found this species common. He 
observed them breeding on July 17. In 1908, near Cordoba, Vera Cruz, Dr. H. 
Gadow found a spawning congress of 45,000 frogs. His account is very vivid. 

"Whilst rambling along the edge of the forest we became conscious of a 
noise, at first resembling the mutter of a distant sawmill; but on our reaching 
the other side of a cluster of trees this sound grew into a roar, like that of 
steam escaping from many engines, mingled with the sharp and piercing 
scream of saws. It came from a meadow containing a shallow pool of rain- 
water. In the wet grass, on its stalks, and on the ground, hopped about hun- 
dreds of large green tree-frogs; nearer the pool they were to be seen in thou- 
sands, and in the water itself there were tens of thousands. Hopping, jumping, 
crawling, sliding, getting hold of each other, or sitting still. Most of them 
were in amplexits, and these couples were quiet, but the solitary males sat on 
their haunches and barked solemnly, with their resounding vocal bags pro- 
truding. Every now and then one was making for a mate, and often there 
were three or four hanging on to each other and rolling over. The din was so 
great that it was with difficulty that we caught the remarks that we shouted, 
although we were standing only a iew feet apart. Each sweep of a butterfly 
net caught at least half-a-dcr/cn frogs. 

"Now the grassy pool, where the frogs were closest, was about 30 yards 
square (900 square yards), rather more than the area of a tennis lawn, and 
each square yard held from 50 to 100 frogs many square yards certainly held 
several hundreds each. At the very lowest computation this gives 45,000 frogs; 
and there was, besides, an outer ring of some five hundred square yards where 
frogs were fairly numerous, say from 5 to 10 to the square yard, mostly spent 
females, but these few thousands we may leave out of the reckoning, to under- 
state rather than overestimate the number. Supposing there were only 20,000 
females, each spawning from 5,000 to 10,000 eggs say only 5,000 the total 
would amount to just 100,000,000 eggs. The spawn literally covered both 
ground and water thickly. But the greatest surprise awaited us on the fol- 
lowing morning, when we went to photograph the scene. There was not a 
single frog left; the water had all evaporated, and the whole place was glazed 
over with dried-up spawn! The prospective chance of millions of little frogs 



3 02 HANDBOOK OF FROGS AND TOADS 

was gone, their expectant parents having been deceived in calculating their 
day of incarnation. That was on the 4th of July, several weeks after the be- 
ginning of the rather fitful rainy season" (H. Gadow, Gen., 1908, pp. 75-76). 

"We found H. bandinii common at Cuatotolapam. Most of the specimens 
were taken during night rains on the banana trees at San Juan. At these times 
they were very noisy. During the clay we found them secreted under boards, 
in the bases of such large leaved plants as the 'elephant ears,' bananas, etc. 
They were observed breeding in a pond near La Laja Creek on July 17" 
(A. G. Ruthven, Gen., 1912, pp. 310-311). 

On the basis ot some material from Mr. Sumichrast (taken at Tehuantepec, 
Mexico) Cope (Gen., i88ob, p. 267) wrote: "Abundant, but only seen in the 
rainy season, when it comes to pools, lagoons, etc. to breed." 

On Dec. 31, 1952, Prof. H. J. Swanson of Edinburg, Tex., sent us three 
specimens, 30, 32, and 32 mm., respectively. They had the reticulated sides, 
dark spot below eye, bar on shoulder insertion, and white stripe across vent 
with black spot below. 

Journal notes: June 18, 1930, Brownsville, Tex. That night, went out the 
Military Road 2 or 3 miles where the road comes opposite a rcsaca which is 
%-% mile from the road. From road heard a chorus new to me. My guess 
was H. baudinii. Pell-mell I started. The resaca looked very foul and dirty 
because of ripe, overripe, and green tomatoes afloat. On little bushes or among 
tomato vines heard these new frogs. The note was a heavy f(ec^ given 5-10 
times with an interesting chuckle following. The first frog heard I never 
could locate, though I was almost on it, in such a tangle of grass and vines. 
Finally located one in a bush where water was waist-deep. It was on a branch 
i foot above water. This one was yellow-green. Then followed another. It 
was brown in color and on the ground where the water was shallow. Tried 
for several more. Whenever I approached, they stopped. Did my white shirt 
or something cl&c give me away? For some time raced back and forth be- 
tween two Hyla buudtmi and three narrow-mouthed toads. Didn't get the 
two Baudm's frogs. After i% hours' effort turned back for fear taximan had 
gone or thought me lost. By pointers (stars) worked my way back. Came out 
by the car. In a pond beside the river heard another H. bandmri. It was deep 
green on back. Only the brown specimen had markings on back. The yel- 
lowish green and deep green specimens were all green on the back. The last 
one was found on the surface of the water among a clump of weeds. 

August 22, 1930. Went over to Mrs. Olive Wiley's museum in the Min- 
neapolis Public Library. She had a beautiful Hyla baudinii from Central 
America. It came on a bunch of bananas to a local merchant. 

Authorities' corner: 

A. Gunther, Gen., 1885-1902, pp. 271-272. R. Kellogg, Gen., 1932, p. 161. 
J. K. Strecker, Jr., Tex., 1915, p. 50. W. G. Lynn, Gen., 1944, p. 189. 



TREE FROGS: HYLIDAE 



303 



Green Tree Frog, Carolina Tree Frog, Marsh Tree Frog, Cinereous Frog, 
Bell Frog, Fried Bacon Frog, Cowbell Frog, Bull Frog 

Hyla cinerea cinerea (Schneider). Plate LXIV; Map 22. 

Range: Virginia to Texas, up the Mississippi River to Illinois and Missouri. 

Habitat: Swampy edges of watercourses; on the taller water plants in 
ditches or pools; on lily pads, trees, bushes, or vines not far from water. 
Prof. E. A. Andrews in 1928 reported finding them in pitcher-plant trumpets. 




Map 22 

The frogs may be on the bushes and stems above the water, but more fre- 
quently are at the water's level. During the day they rest mainly on the stems 
above the water. They may be hidden in clumps of Spanish moss or under 
flakes of bark on trees. June 27, 1922, at Camp Pmckncy, we took Hyla cinerea 
in Pond No. 2, on a vertical wild rice stem, the frog some 3 feet above the 
water. July 5, at Pond No. 3, on the wide-leaved grasshkc plants were some 
7 or 8 Hyla cinerea perched 1-3 feet above the water. On August u, found 
3 males, 2 feet above the water. 

"The green tree frog has been found climbing out on the emergent portions 
of water hyacinths on several occasions. Concerning this species, Kilby (MS 
1938) says: 'By far the greatest concentration of individuals is found among 



304 HANDBOOK OF FROGS AND TOADS 

lake and pond shore vegetation and on the floating rafts of water hyacinths 
that are common in Florida Lakes.' My own records are for January, Febru- 
ary, and October" (C. J. Coin, Fla., 1943, pp. 148-149). 

"This is the most beautiful tree-frog of our fauna. It lives on the leaves of 
plants, frequenting especially lily pads and other aquatic vegetation at the 
edges of lakes. It occurs also, at times, in fields of corn. Its food consists of 
insects, the common fly being, it is said, preferred. Its note resembles the tone 
of a cow bell heard at a distance. Where abundant about water, the frogs are 
very noisy just before dusk, the chorus being broken, however, by longer or 
shorter intervals of silence. A single note is first heard, and as if that were a 
signal, it is taken up and repeated by a dozen noisy throats till the air is reso- 
nant with the sound. After a time it ceases as suddenly as it began, to be again 
resumed after a period of quiet" (H. Garman, 111., 1892, pp. 347-348). 

Size: Adults, i^-a 1 /^ inches. Males, 37-59 mm. Females, 41-63 mm. 

General appearance: This is a very slim, smooth, bright green tree frog with 
a light side stripe, pointed head, and shallow face. It is relatively the longest- 
legged Hyla of the East. Often there are small gold or yellow spots on the 
back. The under surfaces are white or yellowish white. 

Deckert (Fla., 1915, p. 3) described it as "an aristocratic looking tree toad, 
with its long, slender figure of the brightest green, edged on each side with 
a band of pale gold or silvery white." 

"This is a larger species than Hyla lateralis, the length of which according 
to Daudin, is *un pouce et demi au plus.' Dr. Holbrook's specimen, however, 
measured i% inches. The largest specimen in the collection of the Academy 
measures i|X> inches (Fr.). It is a much more slender animal than semifas- 
ciata. In laterahs (viridis, Holl.) the lateral stripe extends as far as the anus, 
and there is a white band running the whole length of the tibia, both ante- 
riorly and posteriorly. The anterior band is absent in semifasciata" (E. Hallo- 
well, Tex., i857b, pp. 307-308). 

Color: Okefinokee Swamp, Ga., April 26, 1921. Male. In dark olive-green 
males there are twenty orange spots scattered over the back. The frog is 
lighter green on sides below lateral stripe, which is cream-colored and ex- 
tends almost to hind legs. Obscure on snout ahead of eye. Cream colored or 
white stripe on back of forearm. Same on back of lower leg and along hind 
end of foot. Throat from angle of mouth to slightly back of chin green. Chin 
proper yellowish cream. Back of these two areas, yellowish cream and green, 
is the wrinkled pink area of the throat proper. One light-colored male has 
stripe straw-colored and extending only an inch behind insertion of arm area. 
There is black above and below the stripe. (Non-Ridgway.) 

Another male. Back may be apple green, dark olive-green, greenish olive, or 
deep slaty olive to almost black, the stripe on side light dull green-yellow or 
clear dull yellow. In one male, stripe not beyond tympanum and suggestive 



TREE FROGS: HYLIDAE 



305 



of H. evittata. The vent, forearm stripe, foot stripe, and heel are white. One 
dark male had the foot stripe fainter. Iris russet-vinaceous. 

Female. Has no pink chin, and green extends in on sides of throat for slight 
distance. Straw-colored stripe to insertion of hind legs prominent, also stripe 
under eye. Under parts from groin to chin the same cream color. (Non- 
Ridgway.) 

Structure: Slender, flat in body; skin smooth or minutely granular; breast 
fold present; vocal sac a round subgulai pouch; a tympanic fold from tympa- 
num to base of arm. 

Voice: The voice is loud and at a distance sounds like a cowbell. The indi- 
vidual call is quon/t, qnonl{, quvn\, qnan^ To some ears, fried bacon, fried 
bacon. To Deckert, grab, grab, grabit, grabit. This is easily one of the most 
characteristic anuran voices of the South. It is one of the rain signals for the 
residents, who call it the "ram frog." In the daytime when the weather is 
sultry or especially in the evenings of late May or early June, some of the 
immense choruses are not easily to be forgotten. Sometimes a chorus starts 
suddenly, quickly reaches its crest, and ends abruptly, to be resumed later 
after a shorter or longer sharply defined interval. Along some of the water- 
courses like Billy's Lake one lone frog near by will begin, then stop, but before 
he has finished, another just ahead of the speeding boat has taken up the task. 
Thus the chorus may travel along the margin of a lake for considerable dis- 
tances. 

At Flatwood, Ala., 1917, after 6:30 P.M. in a drying-up, swampy pond, we 
heard a chorus which sounded to one member of the party like an exhaust 
running into an oil-well pipe. We took one frog with the aid of a flashlight. 
It was on some bushes. Its sides and throat looked like a pink ball. 

The calls may be given about 75 to the minute. The frog may redouble its 
speed and at the same time add a rolling quality to the note: crron\ crron^ 
crronf{. Two individuals calling together but not in unison produce bo babe, 
bo babe, bo, babe. 

J. Le Conte (Gen., 1856, p. 428) said it was commonly known as the bell 
frog, its notes resembling the sound of small bells. 

J. E. Holbrook (Gen., 1842, IV, 120-121) wrote: "Their noise proceeds from 
a single note, which at a little distance is not unlike the sound of a small bell; 
and there seems in general to be one leader of their orchestra, and when he 
raises his note, hundreds take it up from all parts of the corn field, and when 
he stops, the concert is at an end, until he again begins." 

Breeding: This species breeds from April 15 to August 15. The black or 
brownish and white or cream eggs are in small packets or films at or near the 
surface, attached to floating vegetation. The outer envelope is poorly defined, 
becoming part of the mass. The egg is Ho-Vio inch (0.8-1.6 mm.), the inner 
envelope V\^r% inch (2.2-3.4 mm.), the outer envelope %-% inch (3.6-4.0 



306 HANDBOOK OF FROGS AND TOADS 

mm.). The tadpole is medium, i% inches (40 mm.), its tail long and acumi- 
nate, its body green with a sulphur or ivory stripe on the side of the head from 
snout to eye. The tooth ridges are %. After 55 to 63 days, the tadpoles trans- 
form from July 2 to October, at V&- 1 1 A o inch ( 12-17 mm.). 

"The breeding season extends from late spring to late summer. . . . The 
condition of the weather is a controlling factor in determining when, within 
this period, breeding actually occurs" (J. D. Kilby, Fla., 1945, p. 82). 

Journal notes: The same conditions do not always produce the same colora- 
tion. In one case, we had many Hyla ctncrca m a botany drum. All were light 
green except three, one of which was almost black, another olive green or dark 
green, and the third yellowish green. One noon in 1917 we stopped in a low 
woods near one of the bays between Pass Christian and Bay St. Louis, Miss. 
On the saw palmetto leaves we found no end of Hyla cmerea and Hyla squi- 
rella in all color phases. These creatures were on the leaves or in the bases 
between two stems. In one place found two specimens, one a brown phase 
and one a green phase. Some have a yellowish or orange color on the posterior 
faces of thighs, others have purplish. Some have yellowish line on lip and a 
few beside* have faint yellowish line on side. May have a purplish area from 
nostril to femur along the side. In one clump one yellowish green on back, 
another dark greenish, and a third purplish brown. There were unspotted 
//. evittiita; Hyla cinerea, olive, green, or brownish with or without lateral 
bands, with or without dark borders to band. 

May io, 1921. This evening at X P.M. (temperature 72) went to new pond. 
. . . Heard a Hyla cinerea near by. Hcgan to search. It stopped. Later thought 
I saw it. Sci/cd it and it proved a ripe female, not croaker. This female was 
in a cypress 4 feet above the water. Went away. Later the croaker started again. 
On the flat side of an ins leal $ feet from the captured female was the male. 

At night many individuals arc seen to be fairly covered with certain tiny 
insects that arc common in the ground vegetation. Some that were collected 
proved to be harmless flics (Osama longgpes). Their perching on the frogs is 
probably accidental. 

June 14, im f Bccville, Tex. All of a sudden several Hyla cinerea began 
croaking around the pond. They were on near-by mesquite bushes or on small 
dead plants above the water. 

June 15, Beeville, Tc\., at night. Acns began first, soon to be joined by 
R. pipiens, then came M, cincrca, and finally one or two Microhyla ohvacea. 
, . . We caught several //. cinerea. In one mesquite or papilionaceous plant 
found t\vo males facing each other, and caught each by putting the light be- 
tween my legs, and grabbing with each hand. //. cinerea were about I or 2 
feet above the \\atcr. A beautiful brown garter snake (Thamnophis eqttes) 
was coursing around the pond, ostensibly for frogs. 

June 17-21, Brownsville-San Benito, Tex. In various rcsacas were several 
choruses of H. ctnerea. 



TREE FROGS: HYLIDAE 



307 








Plate LXIV. Hylacinerea cinerea (X%). 
3. Females. 2. Male. 



Plate LXV. Hyla cinerea cvittata. 1,3. 
Males (Xi%). 2, Female (XO- 



308 HANDBOOK OF FROGS AND TOADS 

Authorities 9 corner: 

W. Bartram, Gen., 1791, p. 277. E. A. Andrews, N.C., 1928, pp. 269-270. 

P. H. Pope, Tex., 1919, pp. 95-96. J. D. Kilby, Fla., 1945, pp. 81-82. 

V. R. Haber, Gen., 1924, pp. 1-32. G. L. Orton, La., 1947, pp. 364-369. 

Miller's Tree Frog, Green Tree Frog 
Hyla cinerea evittata (Miller). Plate LXV; Map 22. 

Range: Virginia to Maryland. 

"The series cvittata in the American Museum differ from these cinerea in 
having a more vertical, less sloping profile to the snout. The former race is 
also said to differ from cinerea m its broader head but our series of ctnerea ex- 
hibits a great variation in width and no constant difference can be found here. 
It thus appears that typical cinerea has a more northern range than Wright 
and Wright (1933) have assumed. It follows that cvittata has a more restricted 
range for we found only cinerea m the Cove Point-Solomon's Island area. It 
would be interesting to know if all green frogs north of the Washington area 
are not referable to cinerea" (G. K. Noble and W. G. Hassler, Md., 1936, 

P-fy) 
Since 1912 we have been seeing evittatc, semifasciate, and striped cinerea 

from Virginia to Bay St. Louis, Miss., and did not assume as intimated above. 
We merely by our action provisionally called the northern population (varia- 
ble though it is even at Mt. Vcrnon where we have collected it) evittata. Col- 
lectors fiom West Virginia to Florida to Louisiana have recorded or collected 
evittatc cinerea. It is interesting to see that E. R. Dunn (Va., 1937, p. 10) so 
views the northern collection of material: 

"To sum up: ST per cent in the upper tidewater Potomac area have no stripe 
or a short stripe; 41 per cent in other parts of Maryland and Virginia have 
no stripe or a short one. Carolina material available to me is not very extensive, 
but it would seem that there only 25 per cent have no stripe or a short stripe, 
whereas 7*5 per cent have a long stripe. Reports from further south indicate 
that TOO per cent long stripe occurs in the far south, especially on the Gulf 
("oast. 

"We arc, therefore, faced with two opposed populations, obviously differ- 
ent. One occurs in the upper tidewater Potomac; the other occurs in the far 
south. An intermediate population occurs over a wide area. Unfortunately a 
somewhat intermediate population, that of the Carolinas, was named first. 
This seems to be nearer that of the far south, so that Hyla cinerea cinerea may 
be properly applied to specimens of Hyla cinerea from the Carolmas south. 
The name Hyla cinerea evittata may be properly applied to the upper tide- 
water Potomac population. The rest of the Maryland and Virginia popu- 
lations are, and should be considered, intermediate between cinerea and 
evittata. 



TREE FROGS: HYLIDAE 309 

"The most northern locality yet known is the western end of the Chesa- 
peake-Delaware canal in Cecil Co., Md., reported to me by Mr. Joseph 
Cadbury. It is unknown from Delaware or from the eastern side of the Del- 
Mar-Va peninsula.'* 

Habitat: These frogs live in lily pools and reed beds of tid^l marshes. At 
other times, they are found on bushes and small trees near the water. 

"Very little is known about the habits of Hyla evittata. In June and July the 
animals are to be found in the rank vegetation of the tide marshes. Here they 
remain quiet during the day, but as evening approaches they become active 
and noisy. Their food at this time consists chiefly of a small beetle that is found 
on the leaves of the pond-lilies. , . . Later in the season the frogs leave the 
low marsh vegetation. As they are then perfectly silent they are difficult to 
find, though occasionally one may be seen in a bush or small tree, but never 
far from water" (G. S. Miller, Jr., D.C., iH w , p. 78). 

Recently (Va., 1944, p. 1187) Dr. P. Bartsch wrote: "I have been rather inter- 
ested in the habits of Hyla evittata Miller, as observed on our farm, which 
joins Pohick Bay, a tributary of the Potomac River, some 21 miles south of 
Washington. Our house is approximately two-fifths of a mile from the water 
at an elevation of 155 feet. 

"In the late summer when the young hylas have reached a little more than 
half their adult size they leave the lily pads and bonnets of the bay and move 
inland. At this time the frogs come to our upland level, where we find them 
particularly partial to okni plants. I have found as many as six attached to the 
stems of a short row of this vegetable; they blend beautifully into the okra 
color scheme. 

"All the specimens caught on our place arc evittata; that is, they lack the 
band characteristic of H. ctnerea. On the north side of the Potomac River at 
Foxes Ferry I have found the striped form predominant and the evittata color 
scheme only slightly represented." 

Size: Adults, I'/j-i^s inches. Males, 36-47 mm. Females, 32-47 mm. 

General appearance: This slim, smooth, green irog is like Hyla cincrea, but 
without the light stripe on the sides and legs. The under parts are white, 
ivory yellow, or maigucritc yellow, with purple-vmaccous to brownish vina- 
ccous on the front of the foicarms, femur, the rear of the tibia, and more or 
less on the underside of the legs. The rear of arms, legs, and hind feet arc mar- 
gined with white to marguerite yellow. Sometimes there arc fine yellow spots 
on the back. 

Color: Male. Upper parts cosse green or lettuce green; sides and legs javel 
green or dull green-yellow; sides of under jaw from angle almost to tip nar- 
rowly bordered with apple green or dull green-yellow; rear of arms, legs, and 
hind feet margined with white to marguerite yellow; front of legs, feet, axilla, 
and somewhat on underside of forearm purple-vinaceous, livid brown, or 
deep brownish vinaceous; under parts including throat white, ivory yellow, 



310 HANDBOOK OF FROGS AND TOADS 

or marguerite yellow. This individual had faint line on upper jaw tip to angle 
of mouth primrose yellow. 

Female. This one has the faintest line on upper jaw. Upper parts oil green; 
oil green on sides of under jaw as prominent as in male; under parts white or 
cartridge buff; front of forearms, femur, rear of tibia, underside of tarsus and 
foot deep brownish vinaceous. Another female has entire under parts of legs 
and most of pectoral region purplish vinaceous; line on upper jaw white; fine 
lemon yellow or light cadmium spots on the back. 

Structure: Broader head and deeper face than Hyla anerea. 

"Hyla cvittata sp. nov. Type adult & (in alcohol) No. 26,291 U.S.N.M., 
collected at Four Mile Run, Alexandria County, Virginia, July 15, 1898, by 
Gcrrit S. Miller, Jr., and Edward A. Prcble. Zonal position. This frog is 
probably confined to the Upper Austral zone. Geographic distribution. 
While the animal is at present known from the marshes of the Potomac River 
near Washington only, it is to be looked for near the coast from Chesapeake 
Bay to Long Island Sound. General characters. Like Hyla cinerea (Daudin) 
but with broader, deeper muzzle and normally unstriped body and legs. 
Color. Entire dorsal surface varying from olivaceous brown through deep 
myrtle-green to pale yellowish grass-green; ventral surface white, irregularly 
tinged with yellow, especially on chin and throat; colors of back and belly 
fading rather abruptly into each other on lower part of sides; skin of under 
surface of limbs unpigmcntcd, transparent; legs and jaws slightly paler on 
sides than above; eye very bright and iridescent, the pupil black, the iris 
golden greenish yellow, thickly dotted with black, back with a few usually 
less than half a do/en inconspicuous minute, yellowish dots. Measurements. 
Type: head and body, 48; hind leg, 6y; femur, 20; tibia, 21; tarsus, it; hind 
foot, 17; humcrus, 8; forearm, 8; front foot, 10; greatest width of head, 14; 
eye to nostril, $.5; distance between nostrils, 3.5. An adult cf from the type 
locality: head and body, s<>; hind leg, 70; femur, 21; tibia, 21; tarsus, ri; hind 
foot, 17; humeius, 8; forearm, 8; front foot, 10; greatest width of head, 14; eye 
to nostril, 4; distance between nostrils, 3" (G. S. Miller, Jr., D.C. and Va., 1899, 
p. 76) i . 

Voice: The call is very like that of Hyla cinerea. 

"The note is like that of Hyla ptcl(eringn in form, but in quality it is com- 
paratively harsh and reedy, with a suggestion of distant guinea-fowl chatter, 
and scarcely a trace of the peculiar freshness so characteristic of the song of 
the smaller species. The song period continues through June and July" (G. S. 
Miller, Jr., Va., 181)9, p. 78). 

Breeding: Similar to H. cinerea. Smallest specimens in National Museum 
February, 1928, were two young taken by A. S. Miller at Quantico, Va., Oct. 
1 5, igoi. They were 22-24 mm - Another of 26 mm. was taken September, 1923, 
at New Alexandria, Va., by Dr, E. T. Wherry. These are 5-15 mm. larger 
than the 11-17 mm. transformces of H. cinerea cinerea. 



TREE FROGS: HYLIDAE 3 n 

Journal notes: In 1917, on May 29, while encamped on the Alexandria 
(Henshaw Sparrow) Heights, we heard Hyla evittata toward the river. In 
1922, in June, we heard them again from Alexandria. One member of the 
party who knew and had collected H. c. evittata thus identified them. The 
calls impressed the authors as Hyla cinerea. On June 3, 1928, Mr. and Mrs. 
Morris Brady took us to one of the best collecting grounds of Hyla c. evittata. 
We walked along the railroad track and picked up one or two specimens of 
Hyla evittata on the ties, as well as a few toads. On the right-hand side of the 
track, Brady took me to the water's edge where we heard a few calling. It cer- 
tainly sounded like Hyla c. cinerea. Captured one more. Luckily for me, both 
sexes are represented. These specimens are evittate, but it would not be hard 
to conceive of some of them developing a complete stripe or a semifasciate con- 
dition. In fact, Mr. Brady informs me that individual frogs may take on these 
different liveries. The life history of this form, according to what he told me, 
is very similar to that of Hyla c. cinerea. We await with expectancy his account 
of this species. 

Authorities 9 corner: 
G. S. Miller, D.C. and Va., 1899, pp. 75- G. S. Myers, N.C., 1924, p. 60. 

78. O. C. Van Hyning, Fla., 1933, p. 4. 

W. P. Hay, D.C., 1902, p. 199. N. D. Richmond and C. J. Coin, Va., 

H. W. Fowler, Md., 1915, pp. 38-39. J 938, p. 303. 

"The material at hand indicates that the population of this frog inhabiting 
the Del-Mar-Va Peninsula is intermediate between the two subspecies, Hyla 
cinerea cinerea (Schneider) and Hyla anerea evittata Miller" (R. Conant, 
Del., 1945, p. 4). 

Peeper, Spring Peeper, Pickering's Tree Frog, Pickering's Tree Toad, Pick- 
ering's Hylodes, Pickering's Hyla, Peeping Frog, Castanet Tree Frog 

Hyla cmcifer crucifer Wied. Plate LXVI; Map 23. 

Range: Gaspe Peninsula to Manitoba and Minnesota south to northeast- 
ern Kansas, Arkansas, Louisiana, Texas, Mississippi, Alabama, Piedmont 
Georgia, South Carolina to New Brunswick. 

Habitat: These frogs live in open lowland marshes, swamps at sources of 
streams whether wooded or open, sphagnous or cattail, in fact any pool, ditch, 
or shallow pond transient or permanent, grassy or muddy. 

Size: Adults, %-iM inches. Males, 18-29 mm. Females, 20-^3 mm. 

General appearance: This is a small frog with an oblique cross on the back. 
A male when first captured in early spring may be liver brown or bay to claret 
brown. The females are usually lighter in color. Both have obscure bands 
across the fore and hind limbs. The male has yellow on the groin; its throat 
is olive. The under parts are light vinaceous-cinnamon to pale yellow. 

Color: M ale. Upper parts deep olive buflf or Isabella color with light brown- 



3 I2 



HANDBOOK OF FROGS AND TOADS 



ish olive on back and obscure bands across fore and hind limbs; primrose yel- 
low on groin; seafoam green or glass green later ad of vent; below vent, natal 
brown or clove brown spot; throat olive-ocher, or aniline yellow in rear and 
citrine near lower jaw rim; pectoral region sulphur yellow or pale chalcedony 




Map 23 

yellow; rest of under parts light vinaceous-cinnamon; upper jaw cinnamon- 
buff; bister stripe in front of eye. A male when first captured out in the sun in 
early spring may be liver brown, chestnut brown, bay, claret brown, or mars 
brown. 



TREE FROGS: HYLIDAE 313 

Female. Upper parts warm buff, cinnamon-buff, cream-buff, chamois. Bar 
between the eyes, cross on fore part of back, oblique bar back of it on left and 
also right, bar on back near vent, bars on forearms and also on tibia buffy 
brown or drab. Stripe in front of eye and across tympanum and spot on vent 
olive brown or hair brown. Venter pale olive-buff, cartridge buff, or cream 
color. Rear of hind legs and front of thigh aniline yellow, olive-ocher. Under- 
side of femur vinaceous-fawn to light grayish vinaceous. Iris vinaceous-cinna- 
mon or onion-skin pink with some dark spots. 

Structure: Muzzle pointed, projecting considerably beyond the lower jaw; 
skin smooth or nearly so; fingers not webbed. 

Voice: Notes are variable, but shrill, clear, high pitched, and can be heard 
at a distance of half a mile. When at the crest of the chorus stage, the din may 
be heard incessantly night and day, though it is most vociferous in the after- 
noon and evening. The male approaching the pond gives a weaker note with 
a more querulous tremor in it. 

"The vocal sac of the male is largely developed in spring; it is of a greenish 
grey color and lies in loose folds outside the muscles of the throat. Inside the 
mouth are two slits or orifices, opening into the sac, one on each side near 
the angle of the jaws. When the frog is about to give voice the whole body is 
inflated, followed by that of the vocal sac which rounds out into a bubble and 
does not collapse with each 'peep'; the degree of inflation evidently governs 
the volume of sound. About sunset I have frequently been aided in discover- 
ing the frogs by the level rays of the sun striking along the edge of pond or 
meadow and reflecting from this moist, inflated vocal sac with a glittering 
light. 

"Unless the day is overcast, or a warm rain is falling, little is to be heard 
from the frogs till about four o'clock in the afternoon when their concerts 
begin, to be continued in mild nights till morning. Considering the size, the 
volume of sound possible from one frog is surprising. As you approach a 
locality where they are in full voice the air seems to grow gradually dense 
with this ear-deafening, all-pervading sound; occasionally the voices fall into 
a regular measure of time, but the effect is usually a medley of shrill sounds, 
a few voices audible above the others by reason of some peculiarity in key, 
or lack of smoothness in utterance. The piping of each individual is long con- 
tinued; the interval between these musical efforts appears to depend on the 
mood of the musician. One does not note the pause of individual voices in 
the general effect, but, however loud and earnest the piping may be, the in- 
troduction of any unusual sound or appearance, even the low quick flight of 
a bird over the water, is almost sure to give alarm and still them for a while; 
the frogs along the edge of the shore commonly settling away out of sight for 
safety among the dead leaves under water, while those having a position on 
the low bushes or reeds merely cling more closely, flattening the body against 
the object on which they are resting. The interval of silence is brief; soon a 



314 HANDBOOK OF FROGS AND TOADS 

frog rises and gives a shrill 'p ee P>* which is immediately answered by dozens 
of voices. The sounds may appear to come from about your feet, but for the 
reasons given, the chances are against seeing the frogs till some movement in 
the water as they rise from their hiding places, arrests the eye, which on per- 
ceiving one, usually discovers more" (M. H. Hinckley, Mass., 1883, pp. 314- 



Breeding: They breed from April i to June 15. The eggs, singly laid, arc 
submerged among fine grass or other plants in matted vegetation, usually near 
the bottom of the pond, and are 800-1000 in number. They are white or 
creamy and black or brownish in color, the jelly firm with well-defined out- 
line; the egg K'5 inch (0.9-1.1 mm.), the envelope %o-/42 inch (1.2-2.0 
mm.). The small tadpole, i% inches (33 mm.), has tooth ridges % or %. 
After 90 to 100 days the tadpoles transform from July i to Aug. i, at %-%6 
inch (9-14 mm.). 

Journal notes: It is interesting to observe how suddenly a chorus will end 
at one's approach, only to be resumed if the intruder remains quiet. At this 
time, the frogs may be among the grassy hummocks along the sides of the 
ponds, or in the shallow pools within the surface film of dead leaves and 
algae. Not infrequently when disturbed, they may be seen leaping on this 
matted carpet before disappearance. In one instance we discovered one in the 
spathe of the skunk cabbage. 

March 29, 1910. On the 10 A.M. trip, noticed a pair of peepers come to the 
surface. Watched them return to the bottom. The male, as always before, was 
almost as dark as the darkest dead leaf of spring (in the pond) ; the female 
was much lighter, about as light as both males and females are at night. The 
female would lay an egg, then walk or drag along 3 or 4 steps, then stop for 
another fertilization and emission; and so they went on some time through 
weeds and matted vegetation without rising to the surface during my observa- 
tions. Probably they came up occasionally, as when I first discovered them. 
Put them in a wet handkerchief and then in a glass jar where the sun's rays 
could not affect them; took them to a crossroads pond. When I planted the 
jar, they were both lighter in color than when captured, the male considerably 
so. As I put the jar in the pond, they broke embrace but resumed later and 
laid eggs later in the day. 

April 7, 1929. Went to Ringwood (Ithaca, N.Y.) for peepers and wood 
frogs. Went about 6 o'clock. As it grew dusky, the dm grew louder. Peepers 
seemed all around us, in ponds and on land, on the ground and on bushes. 
The calls were everywhere, but to find one and see him call was a real job. 
We both looked for some time without success. Then we both concentrated 
on one small pond near the road that had cattails in it, also a fence post, a 
fallen log or rail, and a few bushes around the edge. It was largely made up 
of tussocks of grass or sedge standing in water. The field grass at one edge 
was wet and "oozy." We knew we must be looking directly at several, but 



TREE FROGS: HYLIDAE 



315 



there was so much noise it was hard to single out one note and locate it. 
Finally I saw one. It was down in the water, just at the surface, standing al- 
most erect, sunken into the edge of a tussock, his back toward the water, the 
throat bubble toward the grass stems. He looked dark greenish brown or 
almost black, like the dirty bases of the grass stems. If his bubble hadn't been 
vibrating, I never would have seen him. At the same instant, Bert caught one. 
Then we found a pair, the female looking much lighter. These were at a 
central tussock. We found another pair at the edge in the "sopping" field 
grass. Then several feet away from the pond, three feet up on a weed stem, 
was a male calling lustily. Then we went over to the ponds in the woods. Here 
were peepers everywhere m ponds and on land. One little fellow was cling- 
ing to a stem a foot or so from the ground, clinging rather diagonally so that 
his bubble went to one side of the stem. We counted his call. It seemed to be 
about one a second, fifteen calls or so, and then a pause. He sat for periods with 
his throat extended. On a log extending into the water, Bert saw six peepers 
calling, but he slipped from the log and they stopped. 

July 14, 1929. Went to Ringwood for adult wood frogs. Along the wood 
road in a fairly open spot where berry bushes and grass are creeping in found 
many transformed peepers. Some were on the ground in an area covered 
sparsely with a fine, soft grass. Many were sitting on the leaves of the berry 
bushes 2 or 3 feet above the ground, others were hopping over the dead leaves 
in the woods. In the pond, along with the numerous newt adults and larvae, 
countless larvae of the spotted salamander, a few transforming pickerel frogs, 
and a very few transforming wood frogs, were many peepers in various stages 
of transformation. 

August 2, 1929. In the same woods, many transformed and a few adult 
peepers were on the forest floor. In the pond, we found no mature peeper tad- 
poles, and only three or four transforming ones. 

Authorities' corner: 
J. A. Allen, Mass., 1868, p. 190. 
M. H. Hinckley, Mass., 1883, pp. 313-317. 
W. I. Sherwood, Conn., 1898, p. 19. 

Florida Peeper, Southern Peeper, Bartramian Peeper, Southern Spring 
Peeper, Sabalian Peeper, Coastal Peeper 

Hyla cruet jer bartramtana Harper. Plate LXVII; Map 23. 

Range: Coastal Plain of southern Georgia and northern Florida. Doubtless 
some of the coastal records of Alabama and Mississippi are of this form. 

Habitat: Similar to that of H. c. cructfer. "Ponds and ditches in and near 
woods; moderately common. Begins to call in December and January, on 
warm nights" (O. C. Van Hymng, Fla., 1933, p. 4). 

"The six specimens from Alachua County, Florida, listed by Harper . , , 



3 i6 



HANDBOOK OF FROGS AND TOADS 




Plate LXVI. Hyla crucijcr crttcifer. 1,2,3. Plate LXV1I. Hyla crucifer bartramiana 

Males (XO- 4- E 8g s (X 1 ^)- 5- Female (XO- 



TREE FROGS: HYLIDAE 

in the original description of this form, were taken in a small woods pond 
which is nearly completely covered with hyacinths. The greatest number of 
individuals I have seen at any one time in northern Florida was in the above 
mentioned pond, where I collected eighteen on the evening of December 31, 
1940" (C. J. Coin, Fla., 1943, p. 149). 

"Northern Florida except in the western portion of the Panhandle. Known 
from the following counties: Jackson, Liberty, Baker, Alachua, Citrus and 
Lake. Habitat. Mesophytic and low hammock, swamp borders, the more 
open bay-heads, and tangles along the smaller streams. Abundance. Moder- 
ately common during breeding season; difficult to collect in numbers during 
summer and fall. Habits. They hibernate and aestivate under logs and bark 
and in knot-holes. In Alachua County they occupy essentially the same habi- 
tats as the much rarer H. v. versicolor" (A. F. Carr, Jr., Fla., 1940, p. 59). 

Size: Harper's type, a male, is 28 mm.; the female topotypes are 31.0 and 
33.5 mm. 

General appearance: "Six specimens of bartramiana from Alachua County, 
Florida, generously presented by Coleman J. Coin, exhibit a still more distinct 
spotting of the under parts than specimens from southern Georgia and proba- 
bly represent the extreme in subspecific characters" (F. Harper, Ga., i93Qb, 

P . 2). 

"Aside from the average difference in spotting, the Coastal Plain specimens 
were noticeably larger than those of the Piedmont" (F. Harper, Ga., 1939!), 

P- 3)- 

Color: Gainesville, Fla., from Coleman J. Coin, Feb. 16, 1946. Descriptions 
of several widely varying patterns follow : 

1. Back orange-cinnamon with bars on the back of snuff brown. Of the 
bars, there are 2 prongs of the cross toward eyes and 2 down sides back of mid- 
body; between lateral bars of cross there may be intermediate series connected 
or not; in front of insertion of hind legs is an oblique bar extending up onto 
the back; also a bar between eyes going onto eyelid; the same color goes from 
nostril through eye and halfway along side. Color of groin, front and rear of 
femur, and lower side of tibia is yellow ocher to buckthorn brown; rear of 
femur heavily spotted amber brown. Upper side of fore and hind legs orange- 
cinnamon with prominent transverse bars on femur and tibia, line on tarsus 
and foot of claret brown bordered above by thread of capucine buff. Venter, 
capucine orange to light ochraceous salmon heavily spotted on breast with 
amber brown; throat light orange-yellow in mid-portion with rim of lower 
jaw black. Iris, lower part snuff brown like vitta, upper part mikado orange. 

2. Back xanthine orange, groin cadmium yellow or raw sienna. No black 
on throat. 

3. Back cinnamon. 

4. Back buffy brown, groin bar goes completely across the back. Rear of 



3i8 HANDBOOK OF FROGS AND TOADS 

femur extensively spotted and bars of tibia extend onto orange area. Center 
of throat olive lake, shading to black on margin of lower jaw. 

5. Back pinkish cinnamon, bars buffy olive. Venter amber yellow, groin 
and rear of femur capucme yellow. 

6. Back Isabella color with face mask and bars of buffy olive; line along 
tarsus madder brown; throat Saccardo's olive with madder brown flecks. 

7. Back mummy brown, no bars showing; bars on legs black; dorsum of 
legs buckthorn brown; orange groin and both sides of femur; line along tar- 
sus claret brown; throat Saccardo's olive to black on margin of jaw; belly 
cinnamon-buff. 

8. Back ochraceous-tawny with Saccardo's olive bars. 

9. Back kaiser brown with markings of carob brown. 

Remarks: March 29, 1946. Live material from Gainesville, Fla., and Ithaca, 
N.Y., compared. Differences most apparent are: the Florida frogs are deeper 
orange on the belly, with more dark specks on under parts and with the back 
of the pecan brown order, whereas those of Ithaca are tawny-olive. When we 
first glanced at the Florida forms we concluded that there were more acces- 
sory arms to the dorsal cross than in northern forms, i.e., 6 or 7 arms instead 
of 4, but some northern ones have the accessory branches also. In general, the 
cross and bars of the Florida form are somewhat broader and more conspicu- 
ous, the bars on legs of some northern forms being very narrow. The belly of 
northern ones is vinaceous, whereas Florida frogs are nearer honey yellow, 
a color strikingly apparent on concealed surfaces of hind legs and underside 
of femur. Furthermore the rear of femur in most of the Florida frogs has 
many fine black specks that are lacking in our Ithaca ones. 

Structure: "Length (snout to vent), 28 mm.; tibia, 14; whole hind foot, 
19.5; elbow to tip of third finger, 13; interolecranal extent (distance between 
elbows when humeri are extended in the same line at right angles to longi- 
tudinal axis of body), 17.5; intergenual extent (distance between knees when 
femora are extended in the &ame line at right angles to longitudinal axis of 
body), 26" (F. Harper, Ga., 1939^ pp. 1-2). 

Voice: "The males sing in buttonbush, briars, willows, Decodon, etc., at the 
water's edge in small ponds, ditches and flooded meadows" (A. F. Carr, Jr., 
Fla., 1940, pp. 59-60). 

(See Psetidacris ornata for Carr's 1933 comparisons of calls of P. ornata and 
H. c. bartranriana.} 

Breeding: "In Charlton County, Georgia, for example, the breeding season, 
as far as may be judged by vigorous choruses, extends from late November 
more or less continuously to early March. Eggs were collected December 9. 
... All (tadpoles) lack the purplish black blotches along the outer edge of 
the tail crests which Wright records . . ." (F. Harper, Ga., 1939, p. 3). 

"January i to March 24. Apparently the season is much earlier in Florida 



TREE FROGS: HYLIDAE 319 

than farther north, where it is April i to June 15 according to Wright. . . . 
I have found them breeding when the temperature was 35 F. P. ornata is the 
only other Florida frog which breeds during such cold weather" (A. F. Carr, 
Fla., 1940, pp. 59-60). 

"In the Gainesville region the peepers usually enter the ponds and marshes 
in late December, and continue to call spasmodically until early spring. My 
only collection records are for February and December" (C. J. Coin, Fla., 
1943, p. 149). 

Journal notes: We have had no experience with this form in the field. 

Authorities' corner: 
A. F. Carr, Jr., Fla., 1940, p. 59. 

Pine Woods Tree Frog, Pine Woods Tree Toad, Pine Tree Toad, Pine Tree 
Frog, Scraper Frog, Femoral Hyla 

Hyla femoraKs Latrcillc. Plate LXVIII; Map 21. 

Range: Coastal region o Maryland to Florida to Louisiana. We rather 
question trans-Mississippian records to Dallas, Tex.; they are possible but 
doubtful. Like Hyla gratiosa, at times H. femoralis betakes itself into the high 
pines and is hard to get, and, also like H. gratiosa, it is a decidedly Sabalian or 
Gulf Strip species. 

Habitat: Trees and shrubs of pine barrens. It breeds in grassy transient pools 
at the roadside or in the woods, in cypress ponds or bays or in lily-covered 
swamp prairies. 

"High pine, high hammock, and all types of flatwoods, occasionally (at 
least) climbing to the tops of the tallest long-leaf pines. ... I have found 
them hibernating in rotten pine logs; one was disinterred at a depth of two 
feet in nearly dry sand, Charlotte County, Dec. 16, 1934" (A. F. Carr, Jr., Fla., 
iQ4ob, p. 60). 

"This species is said to stay high up in pine trees but the only two I have 
personally taken were respectively seven feet up in a small deciduous tree, 
quite likely a sweet gum, near a stream in the first case, and among rank vege- 
tation in the pine barrens in the second case" (C. S. Brimley, N.C., 1940, p. 22). 

R. F. Deckert in 1915 (Fla., p. 3) said: "Hyla femoralis Latreille is called 
the Pine Tree toad, from its habit of frequenting the tops of pine trees almost 
exclusively during the summer months." 

"In 1925 Viosca in discussing the Florida Parishes or uplands of Louisiana 
comments that To the east of the Shortleaf Pine area are the Longleaf Pine 
Hills with gentle slopes, and intertwined by winding creeks. Here we have a 
limited representation of practically all species found in the uplands generally 
and in addition, some species which may be said to be characteristic. Hyla 
femoralis is the typical tree frog of this section.' 



320 HANDBOOK OF FROGS AND TOADS 

"At Leroy, Ala., June 12, 1917, Dr. H. H. Knight while sweeping bushes 
and lower branches of trees with an insect net for Capsids caught two young 
Hyla femorahs" (A. H. Wright, Ga., 1931, p. 275). 

Size: Adults, 1-1% inches. Males, 24-37 mm. Females, 23-40 mm. 

General appearance: This frog is commonly a deep reddish brown in color, 
but may be gray or greenish gray. It resembles a small common tree toad but 
is more slender and the black markings do not form a regular cross. There are 
orange or grayish white spots on the rear of the thighs. The under parts are 
white. The throat may be dark. The upper surface has occasional granula- 
tions, the under parts are areolate, granulate on the throat. 

"Cope (1889, p. 371) writes: 'Body short, rather broad, and the entire ap- 
pearance as to pattern of color and shape not very dissimilar from Hyla verst- 
color, from which, however, it is readily distinguishable by the femoral yel- 
low spots; the dark postocular vitta, the absence of light spots under the eyes.' 

"Brimley (1907, p. 1*58) characterizes it as follows: 'Back . . . markings do 
not form an X-shapcd mark. Back of thigh with yellow spots or variegations. 
No light spot below eye. No yellow spots on sides.' 

"Deckcrt (1915, p. 3) who has studied southeastern frogs more intimately 
than most observers says this species 'resembles our own gray tree frog, with 
its rough skm and star-shaped dark patch on the back, but is smaller and more 
slender/ 

"We would consider this species a small species. Cope considers 35 mm. 
above the average size and his largest is 39 mm. Le Come a century ago gave 
1 1 /-i % inches as the adult range and it has been repeated for this little under- 
stood form. It may reach 45 mm., as he says, but of the 140 specimens from 20 
mm. upwards we have none over 40 mm., the average, 30 mm." (A. H. 
Wright, Ga., 1931,^273). 

"The young newly transformed frogs arc often green and both in this color 
phase and when brown arc liable to be mistaken for Hyla sqtiirella from 
which species they can always be distinguished by the yellow spots on the 
back of the thighs" (C. S. Brimley, N.C., 1940, pp. 22-23). 

Color: June 5, 1921. Male. Cephalic half of pectoral fold pale vinaceous- 
drab to darkish grayish brown; posterior half like the belly. Throat darker 
than the belly. 

Female. General color sorghum brown, deep brownish drab, or mars brown 
on back. Black spot between eyes. Another with four points, two behind and 
two ahead, the cephalic ends above tympanum. Two on either side of the 
middle of the back and one over the crupper. Narrow black or deep brown 
line from snout through nostril to eye, and through tympanum to groin, 
where it breaks up into spots. Another of the same color on back of forefoot, 
forearm, front of foreleg, and over vent. Below white line of the vent is a 
black one. Spots on rear of thighs orange to orange chrome or light cadmium. 
One female with grayish white instead of orange spots. Under parts white 



TREE FROGS: HYLIDAE 321 

Pectoral fold pure white to angle of the mouth. Chin white with fine black 
spots. Iris ecru-drab, drab-gray, or pale vinaceous-drab with reticulation of 
black. 

Structure: Upper surface with occasional scattered granulations; belly, un- 
der surface of thigh, and breast strongly areolate; throat granulate; pectoral 
fold smooth or slightly granulate. Tympanic fold usually present. Muzzle 
rounded. 

Holbrook (Gen., 1842, IV, 128) alluded to Dumeril and Bibron's mistake 
as follows: "Dumeril and Bibron consider this animal as identical with the 
Hyla sqitirella, from which it is, however, perfectly distinct." 

In 1856 John Le Conte (Gen., 1856, p. 428) referred to the same matter. "It 
is wrong in Dumeril and Bibron to say that this species is a variety of Hyla 
squirella. In shape and size the difference is not considerable. The latter ani- 
mal during the warm season is always to be met with about the houses, the 
Hyla femorahs never. Besides, their notes are entirely different." As late as 
1882 Dumeril and Bibron's mistake somewhat influenced Boulenger (Gen., 
1882, p. 398) in his statement, "Appears to be specifically distinct from H. squi- 
rella." 

Voice: A peculiar cicada note in chorus, the chorus one continuous stridu- 
lating dm going down the piny woods like a wave. When high in the trees, 
this frog calls ^<?^ at intervals. When rain brings the frogs close to the water 
and the evening congress is on, the calls are speeded up. There may be as 
many as 6-7 a second, with 60-70 calls in rapid succession without deflation of 
the throat. 

"Le Conte, 1856, seems to be the first to note that it differed from H. squi- 
rella in Voice. No one since has remarked about its voice until Deckert (Fla., 
I 9 I 5> PP- 3"4) wrote, The noise resulting from the calls of the males on these 
occasions, is deafening. This call cannot be reproduced on paper, being a rapid 
succession of harsh, rattling notes higher in pitch than the call of H. sqitirella, 
and kept up all night. During the dry season this tree toad occasionally calls 
from the tops ot the pine trees, one answering the other.' 

"In 1921 the first general calling of Hyla femoralis came May 14. Then the 
author went to Billy's Lake Landing and worked eastward. Heard plenty of 
Hyla jemorahs in trees. They are approaching or are on the edge of cypress 
ponds. Some are yet high in the trees. They are calling at intervals all over the 
piny woods, particularly near the edges of cypress bays. One call, an interval, 
one very faint call, interval. 

"On May 16, when a threatening storm passed over, Hyla femoralis from 
the trees were almost in chorus. Around cypress ponds several calling. One 
was on a projecting piece of pine bark on the tree and within reach, about six 
feet from the ground. Harper saw it croak. Afterwards the throat pulsated all 
the time. Harper notes that it 'let me come up in plain sight, within a couple 
of yards and croaked for me on its bark slab-perch. Throat kept distended 



322 HANDBOOK OF FROGS AND TOADS 



while its sides, or rather whole body, vibrated. Ke\ 9 J(e^ t about 20 times, 
usually ending in %ra\, \ra^! Later in the evening we visited this croaker but 
it was gone. Soon we understood. In two or three trees low down we heard 
Hyla femoralis. Their notes are speeded up, more extended and continuous 
than when high in the pines. One we found on the moss. The instant we en- 
tered another pond we heard a queer note and it was from the trunk of a bay 
tree right near us. It proved to be a male Hyla femoralis. All around us they 
were calling. One was in a bush three feet above ground. Another on moist 
ground at edge of pond in amongst six-foot sedges. When a male croaks it is 
the lower throat which swells out; the chin region does not. Brought back 
three. . . . 

"During a congress when several species are breeding in the same pond the 
machine-gun calls of Hyla jemoralis make it difficult to hear or time other 
calls. Once when we were timing the intervals in Mtcrohyla's calls, Hyla fe- 
moralis calls broke in frequently. In 1921 and 1922 we recorded several in- 
stances where their calls drowned out those of Hyla gratiosa, Hyla squirdla, 
Bitfo quercicus, and Psetidacris oculans. We tried the experiment of half clos- 
ing our ears to shut out the Hyla femoralts calls. The other sounds came out 
very distinctly in the attempt" (A. H. Wright, Ga., 1932, pp. 277-278). 

Breeding: This species breeds from April 20 to Sept. i. The eggs are in 
groups of small films on the surface or just below it attached to grass blades 
of floating roots. The jelly is loose and sticky, the eggs arc brown and yellow- 
ish, theii size Ko-^n inch (0.8-0.9 mm.), the inner envelope Via-Viz inch 
(1.4-2.0 mm.), the outer envelope not distinct, %-% inch (4-6 or 8 mm.). 
The eggs hatch in 3 days. The tadpole is small, i% inches (33 mm.), its tail 
tip acuminate and free of spots, the lower musculature with a light stripe. 
Many have bright red in their tails. The tooth ridges are %. After 50 to 75 
days, the tadpoles transform from June 16 to October, at }> inch (13 mm.). 

"We found breeding congresses in grassy transient pools near the roads in 
the piny woods, in open ponds in cutovcr roads, in pools or ditches beside the 
railroad and roads, in cypress ponds and in cypress bays. We found mated 
pairs in overflowed grassy fields, shallow transient depressions, in temporary 
overflows or drenched cultivated fields, swamps or dreens in the cypress bays. 
Tadpoles were taken in pools beside Indian mounds, railroads, roads, in cy- 
press ponds and sometimes on the prairies, in diverse ponds on the east main- 
land or in bays outside the swamp" (A. H. Wright, Ga., 1932, p. 284). 

"Dangers. This species lays in any pine barrens pool. Many a shallow grassy 
pool will have packets of their eggs. Many are caught by hatching; certainly 
many of the tadpoles never mature. There must be a great loss in this species. 
On July 3, 1921, they were laying in the flooded furrows of cane, corn and 
sweet potato fields. There all were lost. On June 4, 1921, we made the notes 
Tonight the frogs of several species (Hyla femoralis included) are laying in 
all sorts of transient places' " (same, p. 287). 



TREE FROGS: HYLIDAE 



3*3 



Journal notes: May 19, 1921, Okefinokee Swamp, Ga. On my way back 
from a trip to Crosby Pond, at 6 P.M. near the remains of an old cypress pond 
in piny woods saw a female Hyla femomlis hopping along into saw palmetto. 
It was as whitish gray as any Hyla versicolor I ever saw. The spot in middle of 
back showed beautifully, also spot between the eyes. . . . This female I took 
out to look at. It leaped away onto the gray sand. Had a hard time seeing it be- 
cause it matched the gray sand so well. In one minute since its capture, it had 
darkened considerably. 

In denser cover 9 inches high with small saw palmetto and small bushes 
found a half-grown Hyla femorahs. It was green on its back (very suggestive 
of H. sqinretta, which strangely enough we don't get here). 

May 21, in the compartment of Hyla femoralis, most of the specimens that 
were green when captured, in fact all (including one little half-grown one), 
are now Vandyke brown or moss brown. Adults are not often green, but their 
life at transformation frequently starts in a green livery. 

A captive female in a jar, June 19, is pale light mouse gray on back with no 
markings. 

On April 23, 1921, the boys found two on a rail fence at 2:30 P.M. The next 
day they brought three more from the same fence. On April 26, the boys 
found some more Hyla femorahs m the rain barrels along the railroad and 
near the lumber company's woodpile. In a pine near camp about 15 or 20 feet 
up on the end fork of a large branch is a Hyla femoralts male. It doubtless is 
the one we have heard ever since we have been here. 

May 1 8, their calls made a perfect dm in cypress bays and north edge of 
Long Pond. Some were m bushes, others in bay trunks, and others were hop- 
ping in moss edges or grassy edges of the pond or among the lizard's-tail. 
Another congress June 4, 1921. On one palmetto within 2 or 3 feet of each 
other were five male H. fenwralis calling. 

July 11, 1922, 7:30-11 P.M. We heard no end of Hyla femoralts in every 
crossing and cypress pond. . . . They were especially centered about a clump 
of saw palmettos. Here Miles Pirnie tound one pair and I another. Later, 
on one blade were two more pairs and one male. In this clump were possibly 
20-^0 males calling. This breeding came after a hard rain. The females doubt- 
less do not enter the ponds until about ready to lay. 

July 6. Mrs. Chesser hoed out another female in the open field. There has 
been a chorus of them m the pond near by. Probably the female was resting 
here during the day. 

Authorities' corner: 
J. E. Holbrook, Gen., 1842, IV, 128. 
R. Dcckert, Fla., 1915, p. 3. 
A. H. Wright, Ga., 1932, pp, 293-294. 

One of the least known forms of the Coastal Plain is Hyla femoralis, which 
we have always associated with cypress ponds, bays, and thickets. A very sig- 



324 HANDBOOK OF PROGS AND TOADS 

nificant extension of its range comes in its recorded occurrence by J. A. Fowler 
and Grace Orton (Md., 1947, p. 6) at Battle Creek, Maryland. 

Barker, Barking Frog, Coat Bet, Florida Tree Frog, Georgia Tree Frog, 
Florida Hyla, Bell Frog, Giant Tree Frog 

Hyla gratiosa Le Conte. Plate LXIX; Map 19. 

Range: North Carolina (B. B. Brandt) to Florida to Louisiana. Hartselle, 
northern Alahama (Cahn). 

Habitat: Trees of hammocks, pine barrens, and bays. Breeds in pine bar- 
ren ponds and cypress ponds. 

"High pine, high hammock, and dry flatwoods, in the upper branches of 
long-leaf and slash pine and of live-oak. . . . Uncommon locally and gener- 
ally. Wright (1953) describes the rain-song; this odd call may be heard rarely 
from the treetops in high pine. On April u, 1933, H. K. Wallace and I dug 
two adults and a yearling out of slightly moist sand more than four feet deep 
in an Indian mound, in cutover pmeland in Lake County. On November 24, 
1935, I found another in the same mound at about the same depth. This 
winter-burrowing may be of significance in explaining the fact that gratiosa 
is almost never seen except during warm weather" (A. F. Carr, Jr., Fla., 1940^ 
p. 60). 

Size: Adults, 2-2% inches. Males, 49-68 mm. Females, 50-68 mm. 

General appearance: This is our largest native tree frog. It is ashen gray, 
purplish, or green in color. The skin is evenly granular, the back evenly cov- 
ered with elliptical or round spots darker than the general color and encircled 
with black. These spots may be absent in some of the color phases this frog 
assumes. A light stripe extends along the sides, bordered below by a purplish 
brown one. There is some yellow on the sides in axilla of arm and in the groin. 
The under parts are creamy or pinkish white. The throat of the male is green 
or yellow with dark spots just back of the chin, while that of the female has 
the center throat light, and the sides of throat and pectoral region sulphur 
yellow. The colored area of throat is encircled with white on the inner side. 
The spots on the side, chin, and rim of jaw are reddish brown. 

Color: "Mule. June 5, 1921. Lettuce green on hind legs and fore legs, oil 
green or cerro green on the back. The encircled spots of the back have a ring 
of mars brown or bone brown or other browns with green ground color 
within. There are occasional spots as big as pin heads or twice so of bright 
green-yellow or greenish yellow or green-yellow. This color is in the groin 
extending to just back of arm insertion where for one half an inch is a clear 
whitish area. The greenish yellow from groin forward demarcates side color 
from pinkish white or creamy belly color. From tip of snout along upper jaw 
under tympanum and along side to within i inch or % inch of leg insertion 
is a pinkish white stripe. Below it is a line of mars violet or taupe brown (pur- 



TREE FROGS: HYLIDAE 



325 






Plate LXV11I. Hyla femoralis. 1,2. Males 
calling (X'/$). 3- Tadpole (Xi%)- 4- Egg 
cluster (XO. 5- Male (Xi%)- 6. Female 
(Xi). 7- Female (Xi%). 



LX/X. Hyla gratiosa, 1,7. Males 
(X%). 2. Male croaking (X#). 3- Male 
in a cornfield at night (X 1 ^)- 4- Female 
(X#). 5- Transforming frog (X%) 6. 
Eggs (X%). 



326 HANDBOOK OF FROGS AND TOADS 

plish brown) which expands behind the angle of the mouth into a large area 
on the side. Same color back of pinkish white or white area back of brachium 
and antebrachium onto the last finger. Same white line around vent with 
mars violet below. Same combination on the knee. Two or three white patches 
on foot with mars violet between and behind them. Back of femur or thigh 
dull Indian purple. No spots. Below the expanded mars violet area of the side 
is a small whitish line %-i inch long, then comes greenish yellow and finally 
the belly color. Just back of the chin is grayish white speckled with taupe 
brown; then lettuce green or oil green from angle of mouth around to angle 
of mouth, the band Va inch wide when not inflated; the wrinkled part light 
orange yellow or deep chrome. This color extends to pectoral fold of skin from 
forearm to forearm. Ins spotted purplish black and bronzy or vinaceous. 

"Males, ... in botany drum. Upon opening the can one was all green 
with a few yellowish spots but not the regular circles, the stripe along the 
jaw and on the side reminding one of the same in Hyla ctnerea. Hardly any 
white or purplish shows anywhere. Another was a vinaceous gray with the 
rings. Another dark green and the others still different. 

"Female. Belly and under side of legs white, a few picric yellow spots on 
side below band, also rear end of white line on side becomes picric yellow in 
the groin. Picric yellow in the axilla and on cither side of light central throat 
and chin region which is white or sulphur yellow with a few spots. Pectoral 
region sulphur yellow. Stripe around snout along upper jaw to groin white 
except for rear and not interrupted as in the male. Spots on back black encir- 
cled. Color of side and chin, spots, rim of jaw, bands on arms hays maroon, 
chocolate, warm sepia or bone brown. Upper parts calliste green, cosse green 
or apple green" (A. H. Wright, Ga., 19^2, p. 302). 

Structure: A large tree frog of heavy build; head broad and short; fingers 
webbed; large disks on fingers and toes; skin has marked secretion giving 
strong persistent odor. Upper parts except arms and legs strongly granulate; 
under surface of hind limbs granulate only on posterior half of thigh, other- 
wise smooth; under surface of arm, prominent breast fold, and lower throat 
smooth or slightly granulate; chin granulate; webbing of hind feet extends to 
the disks which are very large, 3 mm., or 0/5 of tympanum; disks of fore- 
feet, 4 mm., or 0.66 of the tympanum; a dermal fold over the ear to the shoul- 
der and continued often as a loose fold along the side of the body halfway 
to the groin; a tarsal fold; vomerine teeth between inner nares; head ahead 
of eye short, obtuse; vocal sac well developed. 

"During the last spring, whilst I was residing in the lower country of 
Georgia, it was my good fortune to meet with three specimens of the animal 
described below. One of them was taken in the water of a pine barren pond, 
another was found in a cavity of a sand pit, and the third upon a tree in the 
forest. 

"This Hyla is remarkable for its size, approaching in this respect to those 



TREE FROGS: HYLIDAE 317 

found in tropical regions. Two of them were of a greenish dusky; the second, 
who had concealed himself in a hole in the sand, was of a bright pea green, 
but in the space of half an hour he changed to the color of the others, thus 
showing a complete possession of the faculty of changing color at will, so 
remarkable in many of the Batrachia. 

"There yet remains undiscovered and undescnbed in Georgia three species 
of this genus, which have as yet eluded my search. The notes of these are re- 
markably distinct from those of others; I may hereafter be fortunate enough 
to obtain them. 

"Coarsely granulate both above and beneath. Color above varying at the 
will of the animal from bright green to cinereous and to greenish dusky, with 
roundish spots or irregular blotches of darker, or speckled with variously 
shaped dots of the same, all of them with some few small yellow irregularly 
disposed spots on the back and sides. Beneath whitish, more or less inclining 
to yellow or orange. Upper lip white, or white varied with green or dusky; 
lower lips sometimes whitish, in others of the color of the back; in some a 
white line extends from the upper lip along the side to the insertion of the 
hind leg, in others the sides are more with rounded spots of darker and no line 
visible. Irides black varied with golden; tympanum copper-colored, a consid- 
erable depression between the nostrils and the eyes. Chin varied with dusky 
or green, with a slight fold at the bottom; transverse space between the arms 
smooth, without any granulations. Arms and legs barred, with darker, yellow- 
ish or reddish on the under side, the former smooth beneath, the latter granu- 
late on the posterior half; the under side of the posterior half of the thighs is 
smooth. Disks of the toes very large. 

"Length of head and body 2.5 inches; humerus .6; antcbrachium .6; hand 
.75; femur 1.2; tibia 1.15; foot 1.6" (J. Le Conte, Ga., 1856, p. 146). 

Voice: The call is woody, deep, a curious tonl^ f ton^, like someone pounding 
on a hollow, heavy barrel or hogshead. The call in the ponds is coat bet. The 
call from trees as it approaches water is a bark. The vocal sac is a large sub- 
gular vocal pouch. 

The first time we heard this species to know it as such was on June 5, 1921. 
After a period of heavy rainfall, we visited in the evening an oak toad pool 
near the cultivated fields of Billy's Island, Ga., and heard this curious call. It 
seemed at first somewhat like the call of Rana clamitans only too many times 
separated, and actually the resemblance is fleeting and slight. There are 55-60 
calls a minute, the throat often remaining partly inflated between calls. 

Breeding: This species breeds from March to August. The eggs are laid 
singly on the bottom of the pond. The single envelope, Mo-% inch (2.3-5.0 
mm.), is loose, glutinous, and indefinite in outline. The vitelline membrane 
appears as an inner envelope, Kc-Ko inch (1.6-2.5 nun.) in diameter, the 
e gg %5-Mfl inch (1.0-1.8 mm.). The greenish tadpole is medium, 2 inches 
(50 mm.) long, and is the largest Hylid tadpole of the eastern United States. 



328 HANDBOOK OF FROGS AND TOADS 

The tail is long, its tip acuminate with a flagellum, and the tooth ridges are 
%. The tadpole period is 40 to 70 days. The tadpoles transform from July to 
October, at %G-% inch (14-20 mm.). 

"Choruses of more than twenty or twenty-five males are not often seen. 
They call from deep water in small permanent ponds, from March 3 to August 
14. In Gainesville the first choruses have always gathered in the power plant 
water aerator, where the water is abnormally warm; here I have also seen 
them feeding on insects at the lighted entrance" (A. F. Carr, Jr., Fla., 1940^ 
p. 61). 

Journal notes: While we were in the Okefinokee Swamp, Ga., we usually 
recorded this species as Barker, Barking Frog, or Coat Bets. The last refers to 
the normal note in the breeding pools, the others to a puzzle that perplexed us 
for two seasons. 

On July 15, 1921, on Chesser Island we heard of Coat Bet frogs, so named 
from the sound of their breeding call. On July 16, during the morning, we 
heard a barking frog in the trees south of camp. That night in a near-by pond 
was an immense chorus of Coat Bets. In 1922 the barkers perplexed us even 
more. 

Not until July 26 did we solve the puzzle. About three miles along the road 
from Chesser Island to Folkston we heard in the evening in a cypress pond 
to the right of the road some Hyla gratwsa, and beyond them, a barker or 
two. We went after the barker, and found one in a small gum 4-5 feet, pos- 
sibly 6 feet up. It is Hyla gratwsa! I saw him do it. There were two more 
barkers besides the one I caught. Several Hyla gratiosa were in the water 
calling normally. Coat Bets and barkers are one. 

June 5, 1921, Okefinokee Swamp. Later we found other croakers. One was 
on the raised ndgc of land beside a cornstalk. It was green with the spots 
usually figured. Another male was lying flat in the water at the edge of a 
weedy ridge. It was absolutely flat and spread out. It was without the spots 
and uniform dull brownish green, like the color of the water. Another was in 
a furrow between two rows of corn. It was spotted and alert. The first one 
rested on ground more or less horizontally; the second in water horizontally. 
And the third more or less diagonally upright. The call can be heard at a long 
distance. Usually the heels are widely separated at croaking and the lower part 
of the body dips somewhat. Found a male Hyla gratiosa beside a grassy bank. 
He was as big and round as the can top of a mason jar. How his throat would 
pu(T out I 

Authorities 9 corner: 
R. F. Deckert, Fla., 1915, pp. 4-5. 
C. J. Coin, Fla., 1938, p. 48. 
A. R. Cahn, Ala., 1939, pp. 52-53. 



TREE FROGS: HYLIDAE 329 

Pacific Tree Frog, Pacific Tree Toad, Pacific Hyla, Wood Frog (Cooper), 

Pacific Coast Tree Toad 

Hyla regilla Baird and Girard. Plate LXX; Map 18. 

Range: Vancouver Island and British Columbia to Lower California; east 
to western Montana (Rodgcrs and Jellison), Idaho, Utah, Nevada, and Flag- 
staff, Ariz. 

Habitat: On the ground, especially about streams, springs, ponds, swamps, 
and other moist places; irrigation ditches. 

"Extends to timber-line in the Sierra Nevada; occurs in all zones below 
Alpine-Arctic. Inhabits damp recesses among rocks and logs; the ground in 
the vicinity of springs, streams and lakes; rank growth of vegetation, espe- 
cially in marshy places, trees in damp forests; and in open country, burrows of 
various animals" (J. Grinnell and C. L. Camp, Calif., 1917, p. 145). 

"This tree-toad is probably the most abundant batrachian in California, 
where it ranges from sea-level to n,6oo feet. It may be found abundantly in 
marsh-lands, lakes, springs, under the bark of trees and in almost any other 
place where there is continued moisture. It does not appear to congregate in 
such large compact masses as does Hyla arenicolor, but may be fo.und in large 
colonies during the breeding season" (J. R. Slevin, B.C., 1928, p. 115). 

"Rio Santo Domingo at the Hamilton Ranch, 300 ft., lat. ^o45 / ; 37249- 
37250; June 2. Aguaito Springs, 15 miles E. Rosario, 1300 ft., lat. 304'; 37251- 
37256; June 9. On the night of June 2 multitudes of Pacific tree toads were 
strung along the sand bars and along the sandy banks of the shallow, widely 
spread, slow-moving Santo Domingo River. Their croaking was the domi- 
nant night sound. When the toads were alarmed, they would swim to and 
try to hide in the masses of slimy, thread-like green algae attached to fallen 
willow branches and other debris. The species was also abundant at night in 
and around the isolated pools at Aguaito Springs. Individuals beside but not 
actually in the water, if disturbed, hopped uphill to dry brush cover rather 
than down into the water" (L. Tevis, Jr., L.C., 1944, P- ?) 

Size: Adults, T-I% inches. Males, 25.5-48.0 mm. Females, 25-47 mm - ^ n 
the whole, the mature females are larger than the males, except occasional un- 
usually large males. 

General appearance: This small, delicate tree toad is somewhat smaller and 
more slender than the canyon tree toad (Hyla arenicolor)^ which like this 
form has the rear of the thighs uniform, not spotted. This species reminds 
the authors of species of Pseudacris. The disks on the fingers and toes are 
larger, however. It is very variable in color, usually with stripes on the back 
and a triangle between the eyes, and also with a stripe along the side of the 
head. 

One male is light brown in color with a dark brown V between the eyes 
and two rows of large dark spots on the back. It has a conspicuous greenish 



330 HANDBOOK OF FROGS AND TOADS 

black line from the nostril to the eye and from the eye through and beyond 
the tympanum. Then there is a broken line of spots to the groin. There is 
orange or yellow in the groin and on the rear of femur and on the foot. The 
arms, legs, and feet are indistinctly barred with dark. The upper jaw is a 
beautiful light pinkish cream color. The throat of this male is dark in color, 
greenish. In the middle of the light belly is a broad, longitudinal bluish area. 

Another male is bright green, with an indistinct triangle between the eyes 
and round dark spots on the back. The dark mask is bordered above by a 
light pinkish cream line and below by the light jaw. The throat is olive, with 
some orange on the center rear portion. Orange-yellow is conspicuous in the 
groin and on the rear of the femur. 

"None of the thirty-one specimens from the vicinity of Lake Como has a 
color pattern that approaches the uniform bold pattern of the Hyla regilla 
from the Great Basin. Most of them more nearly resemble specimens from 
Washington, Oregon and California. This evidence tends to confirm what 
might be expected, namely, that the population of the Bitterroot Valley is 
related to that of the Pacific Coast through past or present connection along 
the Columbia River drainage. Fifteen of the specimens have a color pattern 
that seems to be peculiar to the Bitterroot Valley in that it is broken up into 
many small units. One specimen, the most extreme in this respect, has 88 small 
dark gray spots of irregular shape on the back. The heavy Y-shaped mark be- 
tween the eyes that is so characteristic of Hyla regtlla is represented on some 
of these specimens only by a narrow branched line. The markings on the legs 
are also broken into many more small units than is typical for the species. 
Since there were more than a thousand specimens in the collection from the 
Pacific Coast area with which our specimens were compared, it seems reason- 
able to assume that all of the major color patterns that might appear in indi- 
viduals from the Pacific Coast were represented, and from this it might fol- 
low that the population around Lake Como is unique in having this finely 
speckled color phase" (T. L. Rodgcrs and W. J. Jellison, Mont., 1942, pp. 
10-11). 

Color: Male. Pullman, Wash., from Dana J. Leffingwell, March 28, 1928. 
Upper parts some variation of dark grayish olive, olive-buff, Isabella color, 
old gold, chamois, sulphine yellow, oil yellow, oil green, green, cinnamon- 
buff, or clay color. Sometimes with brown band through the eye alone, usually 
with bandhke spots of Saccardo's umber, sepia, or brownish olive on the back 
forming a median stripe and one on either side, also with triangle between 
the eyes, and also stripe in front of eye. Some of small spots on sides are at 
times almost black. Stripe from below nostril along upper lip to shoulder in- 
sertion is primrose yellow. Outside the long dorsolateral spot, back is honey 
yellow becoming cream-buflf or cartridge buff on sides. Upper parts of fore 
limbs chamois or old gold, with light brownish olive crossbars. Underside of 



TREE FROGS: HYLIDAE 



33' 



arms olive-ocher, dark olive-buff. A buff-olive line along outer edge of fore- 
arm. Groin aniline yellow as also on front and underside of femur and to 
some extent on underside of tibia and hind foot. Lower part of rear of femur 
aniline yellow with upper edge of medal bronze. There is also a dark edge to 
front of tibia. Belly either side of meson vmaceous-cmnamon or pinkish buff; 
wash of same on underside of fore and hind limbs. Vocal sac when inflated is 
old gold or ohve-ocher; deflated it is buffy olive or olive in front and olive- 
ocher behind. Iris: pupil rim behind and in front broken by black; above the 
rim, lemon yellow; rest of iris, raw sienna or antique brown. 

Female. San Diego, Calif., from L. M. Klauber, March 28, 1928. May be 
like male, but with little yellow on the groin. Front of femur and underside 
of tibia olive-ocher or aniline yellow. Entire under parts including throat 
may be cartridge buff, or throat may be light with indistinct dots of olive. 
Under parts of hind legs avellaneous or wood brown like males. 

Another specimen. Las Vegas, Nev., Aug. 22, 1925. One half-grown frog 
was light paris green on upper parts; no dark spots except vitta; a little of clay 
color or cinnamon buff in middle of back; the fore and hind legs mainly light 
pinkish cinnamon. 

Structure: Upper parts smooth, not warty as is the usual condition of 
H. arenicolor; prominent breast fold; tympanum round. 

Original descriptions under different names: "Hyla regtlla, B. & G. This 
is a species of medium size; the largest individual observed measuring one 
inch and a half from the nose to the posterior extremity of the body, the head 
itself occupying about half of this length. The hind legs are long and slender, 
the web extending only to half the length of the longest toe; fingers compara- 
tively long. The general color is green above, turning to orange yellow along 
the sides of the head, abdomen and legs. Two oblong, brownish black spots 
exist on the occiput, from which two vittae (one pair) of the same black color 
extend along the dorsal region; a similar band passes from the tip of the nose, 
across the eye and tympanum, and along the abdomen, when it is interrupted 
and forms a series ot black and irregular small spots. In the immature state, 
green is the prevailing color; a few black spots being present along the whitish 
abdomen. Specimens of this species were collected on Sacramento River, in 
Oregon and Puget Sound. Drawings from life were made on the spot by 
Mr. Drayton" (S. F. Baird and C. Girard, Ore., 1854, p. 174). 

"Litoria occidental}*, B. and G. Throat smooth. Abdomen, sides of body 
and lower surface of thighs granulated. Tympanum very small. Fingers al- 
most or entirely free; toes slightly webbed at the base; extremities of both not 
dilated. Color above pale chestnut, with obscure or obsolete blotches of darker. 
Beneath white. A few crossbands on the outside of the legs. A dark chestnut 
line beginning at the nostril, passes back through the eye, behind which it 
widens so as to include the tympanum, stopping just above the insertion of 



332 HANDBOOK OF FROGS AND TOADS 

the arm. One or two oblique blotches of dark chestnut on each side. Body 
i Vic inches long; hind leg extended iVo inch. Hab. San Francisco" (S. F. 
Baird and C. Girard, Calif., 1853, p. 301). 

"Hyla regtlla, B. and G. 3 Proc. Acad. Nat. Sc., Philad., VI. 1852, 174. Syn. 
Hyla scaptdaris, Hallow., Proc. Acad. Nat. Sc. Philad., VI. 1852, 183" (S. F. 
Baird and C. Girard, Calif., 1853, p. 301). 

"Hyla scapularis. Sp. char. Head small; body small and slender, olive green 
above, with numerous bluish blotches; a bluish vitta running from the eye 
over the shoulder; total length one and a half inches (Fr.). Description. The 
head is short and small, depressed; the snout somewhat rounded; the nostrils 
are small and circular, looking upwaid and outward, about a line apart, situ- 
ated immediately below the ridge running from the extremity of the snout 
to the anterior canthus of the eye; they are nearer the extremity of the snout 
than the eye; mouth quite large; the tongue is heart-shaped, quite free behind, 
notched upon its posterior border; there are two series of palatine teeth be- 
tween the nostrils, and separated from each other by a narrow intermediate 
space; the eyes are round and project considerably; the tympanum is small 
and circular; the body is flattened, rather slender, much contracted posteriorly; 
extremities slender; the upper surface of the body and extremities present 
numerous small granulations; abdomen and under surface of extremities much 
granulated; the granulations upon the abdomen vary in size, and are closely 
in juxtaposition; chin and throat granulated. Color. Ground color above 
greenish olive, presenting numerous irregular bluish blotches upon the sur- 
face; several deeper colored blotches upon the sides; a bluish vitta, about two- 
thirds of a line in breadth, extends from the posterior part of the eye along the 
sides of the neck over the shoulder, a short distance beyond which it termi- 
nates; upper surface of extremities marked with bluish spots. Dimensions. 
Length of head 5 lines; greatest breadth 5 lines; length of body i inch; length 
of humerus 4 lines; of forearm 3% lines; of hand to extremity of longest finger 
5 lines; length of thigh 7 lines; of leg 8 lines; of foot to extremity of longest 
toe 7% lines; total length i inch 5 lines. Habitat. Oregon Territory. Presented 
to the Academy by Dr. Shumard" (E. Hallowell, Ore., 1854, p. 183). 

Voice: The call is fy-ec^-e^ in rapid sequence. 

"This is the common frog of the coastal areas, whose loud croaking is to be 
heard nightly in the spring about every puddle, pond, and creek. The noise 
made is surprising for so small a creature. It has the capacity to change its 
color considerably and may be bright green, gray or brown in almost any 
shade" (L. M. Klauber, Calif., 1934, P- ?) 

"In the spawning season, or more properly, as long as there is any likelihood 
of spawning, the 'chorus note,' the note which serves to bring the scattered 
individuals together for breeding purposes, is uttered. This, to human ears, 
sounds like kreck-ek and is uttered over and over again in rapid sequence. 
... At any season of the year this hyla may be heard to utter a single pro- 



TREE FROGS: HYLIDAE 



333 



longed note, kr-r-r-eck, lower in pitch than the song note. This note may be 
used, as indicated above, at the ending of a 'chorus'; but it is also uttered in 
other seasons of the year when the individuals are hidden in their daytime 
retreats, and when no 'songs' are to be heard" (T. I. Storer, Calif., 1925, pp. 
221-222). 

Breeding: This frog breeds from early January to mid-May. (See "Journal 
Notes" for 1942.) The brown and yellowish eggs, in small, loose, irregular 
masses (10-70 eggs) are laid in quiet water beneath or sometimes at the sur- 
face, attached to vegetation. They remind one of Pseudacris masses. The egg 
is %o inch (1.3 mm.), the inner envelope (firm) VJo inch (2 mm.), the outer 
envelope (loose, sticky) %<\-% inch (4.7-6.7 mm.). The tadpole is medium, 
i% inches (46.6 mm.), full and deep-bodied, its tail tip acute or obtuse with- 
out a flagellum. It is dark brownish in color and the tooth ridges are %. After 
50 to 80 days the tadpoles transform from May 15 to September i, at ^o- 1 / y ie 
inch (11-17 mm ')' Mrs. Katherine Van Winkle Palmer took ten transform- 
ing Hyla regilla at Oakvillc, Washington, July, 1923. They ranged from 12 
to 15 mm. Our series of 36 from Alta Meadows, Sequoia National Park, Calif,, 
Aug. 22-27, 1917, ranged from 11.5-17.0 mm.; mode 13.5, average 14.5 mm. 

"The egg clusters of this hvla contain a varying number of eggs. In labora- 
tory aquaria and occasionally in the field single eggs are laid. Some masses 
total over 60 eggs. Probably the number in a mass reflects in a general way 
the degree of solitude experienced by a pair when eggs are deposited. Where 
only one female is laying a few large clumps are to be found. Where many 
hylas have been spawning in one place the egg masses are small. 

"At times very large numbers of eggs are laid in a relatively small area. In 
a small pool near the main Thornhill pond I found, on Jan. 25, 1913, 59 masses 
in an area approximately 10 X 15 feet. Additional masses were laid in this 
same area on subsequent nights. Other areas in this same pond which seemed 
to be of exactly the same character were not utilized by the species. 

"The total number of eggs deposited by a single female has been ascer- 
tained in two cases. A mated pair was taken from the Thornhill pond on 
January 31, 1914, and confined in an aquarium. Next morning a total of 730 
eggs was found. The average number of eggs in a clump was 18. Another cap- 
tive which began laying on the night of Mar. 7, 1922, and continued up until 
4 P.M. of the day following deposited a total of 1250 eggs; the average of the 
58 masses laid was between 21 and 22. The largest mass contained 60 eggs and 
there were two with the minimum of 9" (T. I. Storer, Calif., 1925, pp. 224- 

225) - 

"Insemination occurs at the moment of egg extrusion. The male brings his 
cloacal aperture close to that of the female, discharges a quantity of trans- 
parent semen, and with a quick, firm extension slides his feet posteriorly over 
the sides and hips of the female, then deftly retracts to his normal position. 
Simultaneous with this foot action, the female extrudes a clutch of eggs into 



334 HANDBOOK OF FROGS AND TOADS 

the cloud of sperm about her cloaca. Some time before releasing an egg mass 
the female often scratches at the surface of the substratum on which the eggs 
are to be deposited. As the eggs are extruded the cloaca is brought into close 
contact with this surface and attachment is thus effected. The female removes 
any eggs which may partially adhere to the cloaca during extrusion by a pre- 
cise flexor-extension reflex of the hind legs. In this the tarsi are applied directly 
to the adhering eggs and a slow extension effects the removal. 

"Deposition of an egg mass is usually followed by a quiet moment during 
which the bodies of the pair become slightly more inflated than normally. 
The intervals between egg deposition, ranging from 2 to 10 minutes or longer, 
are spent in bursts of vigorous activity, mainly on the part of the female. What 
function this may serve, if any, is conjectural, but it apparently takes place 
under natural as well as artificial conditions. In the laboratory such periods 
are followed by a rest and then further oviposition. Shortly after laying is 
complete, the pair becomes separated. The entire time in amplexus has been 
observed to range from 8 to 40 hours or moie. 

''The total number of eggs deposited was found to range from 500 to 750; 
however, Storer reports an instance of 1250 eggs being laid. The number of 
eggs per clutch is usually about 16, but varies from 5 to 60" (R. E. Smith, Calif., 
1940, pp. 379- ?8o). 

"Several of these frogs were found at Big Lake, together with the north- 
western toads. Egg clusters in the litter on the bottom, near the edge of the 
lake, were in jelly-like masses a little more than one inch in diameter. The 
egg diameter might be from %a to Vf inch" (F. G. Evenden, Jr., Ore., 1945, 
p. 252). 

"Hyla regilla also spawns abundantly in the pool on Alpine Road, the onset 
of breeding seeming to coincide closely with that of A. t. caltforntense" (V. C. 
Twitty, Calif., 1941, p. 2). 

"One male, Scaphiopus /;. intermontantts (amplexus inguinal) when col- 
lected was clasping a female Hyla regilla which died soon after, apparently 
from rupture of the abdominal wall" (G. C. Carl, B.C., 1942, p. 129). 

Journal notes: Aug. 12, 1917. We camped at Jacumba, Calif., beside the 
Mexican border. ... In a little side stream of the creek we found a series 
of H. regilla from tadpole to transformation. In the same place among the 
weeds of the moist area we took six to ten adults very variable in color. 

Aug. 25. In Alta Meadows, we found no end of transformed Hyla regilla. 
In the bog-ten aced pools were plenty of tadpoles and advanced stages. On 
the trail, R, C. Shannon found a full-grown frog and the boys reported a 
large one from Alta Peak. The transformed frogs were in the meadow 
land. 

Aug. 20-22, 1925, Las Vegas, Nev., Tonapah Road. We followed one tiny 
stream back into the field and returned with the trophies of the search: 6 Rana 
fisheri, 4 Hyla regilla, i Bttjo compactilis. In a broad springy area sedgy and 



TREE FROGS: HYLIDAE 335 

shady, we picked up a H. regilla. In one very small sedgy area, we caught four, 
three of which had triangles between the eyes and some dorsal stripes. A half 
grown one was green with no triangle and only the vitta back of the eyes. 
Later at the big springs, we caught one that looked very yellow with very in- 
distinct pattern. We caught several tadpoles. 

Starting in January, 1942, we traveled northward west of the mountains 
from southern California to Washington and southward east of the moun- 
tains, arriving in Nevada in mid-May. The Journal notes of that period give 
a specimen cross section of the activities of this widespread species. 

Jan. 31, 1942. Stayed at Julian, Calif., for night. Last night heard a chorus 
of Hyla regilla. The early season and the chorus each make me think of 
Pseudacris choruses in early spring. It is so unlike most true Hyla barks of 
the U.S.A. 

Feb. i. Went to Cuyamaca. In Julian region where I heard the choruses last 
night took about seven or eight male H. regilla. One female and one very 
small one. Tonight the male is mated axillary fashion. Tonight heard a large 
chorus of H. regilla but no others. 

Feb. 7. Five miles out of Monterey at 12:30 P.M., 7oF., sunlight. We heard 
chorus of H. regilla in an overflow grassy area near base of pine hill. 

Feb. 8. Salinas to Castroville, Calif. Could find no Hyla regilla eggs. Do 
they lay them in dense vegetation at the edge of water like P. feriarum of Car- 
lisle, Pa.? 

Feb. 12. Visited W. M. Ingram. He had two Hyla regilla masses of eggs 
ready to hatch. 

Feb. 14. Trip to R.mcho del Oso Prof. Theodore S. Hoover owner. Found 
H. regilla in swampy area. Up the creek to sawmill. Here in a little pool found 
very young H. regilla tadpoles. Rodgers took a large H. regilla with reddish 
down middle of back. 

March 8. Stanford artificial lake. Found in an overflow shallow some eight 
fresh masses oi H. regilla eggs. The yellowish underside is evident. Mass an 
inch or less in diameter. Sometimes two masses on one grass blade. One or 
two masses hatching. Found some tadpoles VL> inch long. 

March 17. Later went north of Crescent City on Route 101 about 5 miles. In 
a pond, countless tads of H. regilla and a few egg masses almost hatching. 

March 18. Took Hyla regilla tads Carpcntcrville, Ore. H. regilla calling at 
midday. 

March 20. Up McKenzie River watershed with Miss Ruth E. Hopson of 
Springfield, Ore. In a pond with fallen logs, no end of H. regilla migrating 
to the pond, mostly females. 1800-1900 elevation. 

March 26. Started from Aberdeen at 9 A.M. Foggy. At Springfield School 
Dist. No. 12 saw, beside road, ponds with pussy willows in them. Here were 
Hyla regilla masses, some just hatching. At Humptulips River crossing, in 
pools, caught a Hyla regilla. Two miles farther took a female. 



336 HANDBOOK OF FROGS AND TOADS 

March 27. Hoh to Borachiel rivers. In meadow heard one lone H. regilla. 
Saw here an egg mass of H. regilla. 

March 28. Went to Olympic Park headquarters. Mr. Macy gave me a permit 
and a map. In their pond saw several fresh egg masses and some about to 
hatch (H. regilla). Just as we turned on to Elwah road, in one meadow pond 
saw some 500-800 egg masses of H. regilla hatching and fresh. Anna caught 
a pair. 

April 4. Just out of Maupin, Ore., to the south of town in a gravel-pit pool 
saw some six or eight H. regilla masses. One H. regilla jumped in. 

April 5. Lapine, Ore., roadside ditch. The H. regilla are calling loudly but 
could find no egg masses. Flatland, open, some cultivated, some meadows, 
trees. 

April 10. From Medford to Project City, in rather barren stream took some 
almost full-grown H. regilla tadpoles. 

April ii. Chico, Calif., in a spring watering trough were large H. regilla 
tadpoles. 

April 12. Out with Tracy and Ruth Storer. Between Michigan Bar and 
Consumne spotted beside road a possible A. t. cahforniense pond. H. regilla 
tads mature. White egret here feeding. 

April 21. Below Tollhouse, Calif., in one pond we took two H. regilla but 
no R. b. sierrae. 

April 23. Turned toward the coast again to Atascadero, Calif. Heard no 
end of H. regilla last night. 

May i. In French Valley, Calif., in drying pool took H. regilla tads, one 
adult. At a culvert took H. regilla tads mature and one transforming. No end 
of Brewers black birds around this pool of tadpoles and aquatic insects. 

May 5. Three and a half to four miles south of Buckman Springs, Calif., at 
bridge below confluence La Posta and Cottonwood Cr., took two H. regilla. 
Saw no H. regilla tads. 

May 14, Springdale, Nev. Bobbie Shoshone went out last night and caught 
one Bufo b. nelsom and several Hyla regilla (3). 

May 15, Las Vegas, Nev, Went out Tonopah road 25 miles and returned 
after dark. Near trailer camp was a large pond with bullfrogs in it and near 
by a big chorus of Hyla regilla. 

Authorities' corner: 

F. C. Test, Gen., 1899, 2I > $* J- Grinnell, J. Dixon, J. M. Linsdale, 

W. P. Taylor, Nev., 1912, p. 343. Calif., 1930, p. 141. 

J. Van Denburgh and J. R. Slevin, L.C., A. and R. D. Svihla, Wash., 1933, p. 

1914, p. 135. 127. 

A. G. Ruthven and Helen T. Gaige, S. G. Jewett, Jr., Ore., 1936, p. 71, 

Nev., 1915, p. 15. 



TREE FROGS: HYLIDAE 



337 




Plate LXX. Hyla regilla. 1,2,6. Females Plate LXXI. Hyla septentrionalis (X%). 

(XO- 34- Mai* (X%)- 5- Female (X%). 1-3- Females. 



338 HANDBOOK OF FROGS AND TOADS 

Giant Tree Frog 

Hyla septentnonalis Boulenger. Plate LXXI; Map 19. 

Range: On Oct. 30, 1931 (Fla., p. 140), Dr. T. Barbour placed it as estab- 
lished at Key West, Fla. In 1914 (p. 347) he had it from New Providence, 
Andros Island, Rum Cay, Cuba, and Grand Cayman. Matecumbe, Fla., and 
farther northward. 

Habitat: Cisterns, wells, outbuildings, drain pipes; banana trees, large leaf 
axils. 

"Old cisterns and damp out-buildings; in the axils of palm and Caladium 
leaves; banana trees; Barbour (1931) records it from drain pipes. Abundance: 
Locally common; I have seen twenty-five or thirty clinging to the walls of a 
single cistern in Key West. Habits: Chiefly nocturnal, but often calling and 
foraging during the day in the dilapidated cisterns which abounded on the 
island before 'New Deal' improvements. During humid weather they may be 
found in trees and shrubs some distance from water. I have discussed septen- 
trionalis with some of Key West's most venerable citizens and have found 
none who can recall a time when the blatant choruses of the 'bull frogs' were 
not to be heard after summer rains. Feeding: They often congregate around 
street lights at night; I have watched them feeding on cockroaches and on 
lepidopterous larvae; they are batracophagous and cannibalistic in captivity" 
(A. F. Carr, Jr., Fla., 1940^ p. 62). 

Size: Adults, 2%-^% inches (64-130 mm.), "There is a great size disparity 
in the breeding males; individuals that I have measured varied between 40 
and 73 mm." (A. F. Carr, Jr., Fla., 1940, p. 62). 

In our Matecumbe Key series (March 18, 1934) we took males 48, 48, 49, 49, 
50, 50, 51, 52, 52.5, 57, 58, 59, 60, and 61 mm. in length, fourteen in all, and one 
small female, 52.5 mm. In 1940 at Key West, Mr. and Mrs. R. H. McCauley, 
May 5, secured two males, 46 and 48 mm., and one large female, 82 mm.; on 
April 10, 1940, they secured on Stock Island, Monroe Co., Fla., two females 
75.5 and 77 mm. in length. The largest we have is from Key West, taken by 
M. K. Brady and O. C. Van Hyning. It is a female, 92 mm. in length. We 
have seen large specimens but recall none over 4 inches. With our limited 
series we can only approximate the sex sizes as follows: males, 46-75 mm.; 
females, 52.5-130.0 mm. 

General appearance: This is a large tree toad. The head is broad; the out- 
line of the skull evident, as the skin is united to the skull; canthus rostralis 
and nostrils very prominent. The top of the head is smooth; the eyelids and 
back are roughened with large and fine tubercles. The most conspicuous char- 
acters are the very large disks on fingers and toes, those on the fingers being 
fully as large as the tympanum. The eyes are large and prominent, the iris 
with brilliant orange tints. The color, when the frog has been under cover, 
becomes a dull olive-green, but in the light becomes citrine, turtle green, or oil 



TREE FROGS: HYLIDAE 



339 



yellow, with indistinct dorsal spots of dull citrine or grayish olive. The legs 
are barred with the same. The rear of the ftmur is reticulated with the same. 
The throat is pale, huffy, and slightly granular; the rest of the venter is con- 
spicuously and roughly granular and dull yellow in color, with the underside 
of the femur a deeper yellow, and the axilla bright yellow with a wash of the 
same color along the sides. The tubercles under the joints of feet and hands 
are prominent and pointed. 

"Male with two external vocal vesicles, each being situated near the angle of 
the mouth; during the breeding-season the inner side of the first finger cov- 
ered with blackish rugosities. From snout to vent 75 millim." (G. A. Bou- 
lenger, Gen., 1882, pp. 368-369). 

Color: Female. Key West, Fla., from M. K. Brady and O. C. Van Hyning, 
May 5, 1932. Trough of casque and concave lorcal region sulphur yellow, ecru- 
olive, or buffy citrine. Postorbital horns of casque pale turtle green or oil 
yellow. Wash of this color i inch or more wide down middle of back. Area 
below and behind eye to below tympanum, upper parts of fore limbs, and 
throat light buff or pale ochraceous-buff. The sides of body with slight wash 
of the same color. From axilla along either side of belly to groin is a wash 
of wax yellow or primuline yellow in axil. Two bars on thighs ceasing halfway 
as they meet reticulations on rear of thigh. Tibia with two complete bars. Bars 
and obscure spots on the rear half of back are dull citrine, grayish olive, or 
buffy citrine. The dark reticulations of the thighs are colored like bars of 
thighs with the interspaces green yellow. There is a little of same color on 
front of femur and in groin. The top of the foot has decided wash of same 
color as top of fore limbs. The digits are turtle green. The center of tympanum 
like back surrounded by ring of mars brown. The iris is capucine yellow or 
ochraceous-buff finely dotted with black. Sometimes the iris has considerable 
of the color of the back or of dorsum of fore limbs. Undersides of fingers and 
toes are cinnamon-drab, 

Structure: Tongue subcircular; head broader than long in adults; casque 
emarginate behind; snout rounded, contained two times in head to tympa- 
num; pollex rudiment not free projecting; disks conspicuous; no interocular 
bar; upper eyelid small; tympanum distinct, %-% diameter of eye. Dunn in 
1937 (Copeia, p. 166) placed H. septentrtonalis in the Hyla bmnnca group. 

"Fingers slightly webbed; no projecting rudiment of pollex; toes two-thirds 
webbed; disks of fingers nearly as large as the tympanum, of toes smaller; 
subarticular tubercles well developed; a slight fold along the inner edge of 
the tarsus. The hind limb being carried forwards along the body, the tibio- 
tarsal articulation reaches between the eye and the tip of the snout . . ." 
(G. A. Boulenger, Gen., 1882, pp. 368-369). 

Voice: "This sound is like the jerky pulling of a rope through an unoiled 
pulley and is very characteristic" (T. Barbour, Fla., 1931, p. 140). 

"The choruses arc unique in their heterogeneity of pitch and timbre, The 



340 HANDBOOK OF FROGS AND TOADS 

call is a rasping snarl somewhat comparable to that of Scaphiopus 
The pitch of non-musical sound is difficult to determine, but I would judge 
that the individual notes vary through at least one octave" (A. F. Carr, Jr., 
Fla., 1940^ pp. 62-63). 

We were first attracted to them at Upper Matecumbe Key by a huge chorus 
near our cabin. In the circle of bushes around a small very deep pond were 
these gaint tree frogs. We neglected to characterize their call, to photograph 
them, or to describe them. Doubtless there were eggs, but Elaphe rosacea was 
our chief interest at the moment. 

Breeding: "J une I 4 to September 16 (this period corresponds suspiciously 
with the academic summer-vacation season; however I looked for them in 
vain during the week of December 18-23, 1932). The eggs are laid in tempo- 
rary drainage ditches, in the flooded basements of wrecked buildings, and in 
cisterns. The young emerge at 15.5-16.5 mm. Before the tail is absorbed, the 
back becomes granular and warty as in the adult; the immature differs strik- 
ingly from the adult in the possession of a dorso-lateral white stripe extending 
from the hind margin of the orbit to the groin. E. Lowe Pierce and I found 
all stages, from newly hatched larvae to breeding adults, in an old cistern, 
June 16, 1934. We found a tremendous number (300-400) of recently emerged 
individuals perched on the twigs of a small thorny bush in a vacant lot, the 
night of June 14, 1934" (A. F. Carr, Jr., Fla., 1940^ p. 62). 

March 18, 1934, Matecumbe Key, Fla. Males have enlarged thumbs; on 
some smaller males the horny, sharp-edged ridge is not so black. In general, it 
is a dark, horny excrescence on inner side and upper side of the the thumb; it 
is semilunar, extending from base of thumb to base of last segment of thumb. 
Outer edge acute and making a sharp acute edge, apparent from ventral side 
of thumb. It's an expanded addition or prepollex on side of the thumb 
(pollex). 

Some of these males are almost uniform, some streaked in groin like 
H. batidmii, some very mottled on back; almost all with leg bands, some nar- 
row, some wider. Several with two prominent stripes extending backward 
from eyes. 

May 5, 1940. Dr. and Mrs. R. H. McCauley took one specimen just beyond 
transformation, 40 mm. in length (with 24-mm. body and remnant of tail 
16 mm.). This specimen has the tail crests and musculature heavily mottled 
with black. They also took the following interesting series: at transformation 
21 mm.; past transformation 25, 26, 28, 30, 30, 32, 33, 36, 37 mm.; male with 
black on thumb 46, 48 mm. 

Key West, Fla. "The largest of fourteen specimens collected was a female 
measuring 92 mm. in length of body, and 210 mm. stretched out. Eggs, tad- 
poles, and recently transformed frogs were also collected" (E. R. Allen and 
R. Slatten, Fla., 1945, p. 25). 

Journal notes: March 18, 1934, Upper Matecumbe Key, Fla. Came to Cari- 



TREE FROGS: HYLIDAE 34 ! 

bee Colony. At night heard a great chorus near the water tank. There were 
countless small and half-grown Hyla septentnonalis in the pond. No end of 
Hyla sqttirdla calling. Are H. squirella the prey of the giant tree frog? From 
limbs higher than my head big tree frogs would leap into the middle of the 
pond with a pronounced ^erplunf^ (like a toad). In looking for the frogs 
Anna espied in the bushes a fine Elaphe rosacea. Does it feed on the frogs ? 

Authorities' corner: 
G. A. Boulenger, Gen., 1882, pp. 368- T. Barbour, Fla., 1914, p. 238. 

369. T. Barbour, Fla., 1931, p. 140. 

L. Stejneger, Fla., 1905, p. 330. C. S. Brimley, N.C., 1940, p. 23. 

Squirrel Tree Frog, Southern Tree Frog, Southern Tree Toad, Scraper Frog, 
Rain Frog, Squirrel Tree Toad, Squirrel Hyla 

Hyla squirella Latreille. Plate LXXII; Map 24. 

Range: Virginia to Texas, and north up the Mississippi basin to Indiana. 
Near Columbia, S.C., Corrmgton (S.C., 1929, p. 66) "found them on the fall 
line but never in good typical piedmont territory." 

Habitat: In and around buildings; about wells; in bushes, trees, or vines; 
in fields and gardens. It breeds in open ponds in the pine barrens, or in shallow 
roadside pools. 

"This species was found on the ground about shady hammock-land on Boca 
Chica Key. I also saw one on a Gumbo Limbo tree on Vaca Key" (H. W. 
Fowler, Fla., 1906, p. 109). 

"Common when breeding; frequents gardens and is often seen in crannies 
about porches of houses" (O. C. Van Hyning, Fla., 1933, p. 4). 

"Shows little discrimination in major habitat selection. It exhibits some 
preference for the more open wooded areas such as high pine and mixed ham- 
mock and for edificarian situations. Abundance. The commonest Florida 
Hyla" (A. F. Carr, Jr., Fla., 1940^ p. 61). 

Size: Adults, %-i% inches. Males, 23- $6 mm. Females, 23-37 mm. 

General appearance: This species is small, delicate, and with smooth skin. 
The head is short, eyes prominent with black pupil and bronzy iris. The back 
is green or brownish in color with at least a partial transverse bar between the 
eyes and with white on the upper lip. Frequently there are rounded spots on 
the back. The rear of femur is not spotted. There is a light line below the eye 
and over the shoulder. There may be a light, irregular line along the side just 
above the belly. 

"The colours of this animal are even more changeable than in any species 
with which I am acquainted. I have seen it pass in a few moments from a light 
green, unspotted and as intense almost as that of Hyla lateralis, to ash colour, 
and to a dull brown with darker spots; the spots also at times taking on dif- 
ferent tints from the general surface. The markings, too, vary exceedingly in 



34* 



HANDBOOK OP FROGS AND TOADS 



different individuals, the white line on the upper lip and the band between 
the orbits alone are constant" (J. E. Holbrook, Gen., 1842, p. 124). 

See M. C. Dickerson, Gen., 1906, pp. 149-150 for a discussion of the wonder- 
ful change of color in this form. For variability of color in Hyla squirella and 
Hyla ctnerea, see our "J ourn d notes" from Pass Christian and Bay St. Louis 
under Hyla cinerea. We have not seen Bryan P, Glass's (Tex., 1946, pp. 101- 
103) specimens of "A New Hyla from South Texas" but suspect he has Hyla 




Map 24 

squirella in one of its variable disguises in his new Hyla flavigula. Of course 
examination of specimens alone can confirm this impression. Hyla squirella 
adult males and females range as low as 23 mm. and as high as 37 mm. Glass's 
specimens may be about the mean of the squirrel tree frog. This species has 
been recorded four times inland from Aransas country and north of the 
Nueces River, Tex. P. H. Pope (Tex., 1919) found H. squirella the most 
abundant Hyla of the Houston area. 

Color: Males. June 22, 1922. Back citrine or buffy citrine in one; on another 
ecru-olive or light yellowish olive; rear back and Lop of hind legs chrysolite 
green to lime green. Under parts of hand, also hind foot, orange-rufous, xan- 
thine orange, to orange chrome; hind limb posterior and anterior faces and 
throat raw sienna or mars yellow; throat of another aniline yellow; in another 
hind leg parts raw sienna. Stripe on upper jaw to body greenish yellow or 



TREE FROGS: HYLIDAE 343 

lemon yellow. Belly cream color or ivory yellow. Tympanum hazel or russet. 
Iris spotted black and army brown fawn in one, deep chrome and antique 
brown in another. Greenish on either side of throat but not clearly defined or 
definite as in Hyla andersonii and Hyla cinerea males. One male held in the 
hand in good light became courge green and another oil green. A small last 
year's H. squirella was light dull green-yellow above, and upper jaw stripe 
light chalcedony yellow or pale chalcedony yellow. Sometimes those in water 
may be greenish, while those on edges or on land may be brownish. 

Females. July 3, 1922. Four females of four different pairs had under parts 
white, with no discolored throats. Throat slightly primrose yellow, reed yel- 
low, or white. Fore limbs, groin, hind limbs before and aft, tibia, and hind 
feet reed yellow to olive-yellow. In one, yellow ocher fore and rear part of 
femur, underside of tibia, and foot and fore foot. The females, unless very 
large, have none of the bright orange-rufous, xanthine orange, or orange 
chrome of the males. 

Structure: Form delicate; skin smooth; canthus rostralis well marked, but 
not sharply edged; throat of male raw sienna or yellow with light area of 
greenish on either side; vocal sac a large hyaline subgular pouch. 

Voice: This call is a harsh trill, regular, mostly continuous, 15 calls in 10 
seconds, but not very loud. It could not be heard a few rods away. One gave 
67 pumps in 45 seconds. "Call rasping, but not very coarsely so; rather ventrilo- 
quistic; flat; short to Italian a; aaaa or waaaaal^' (A. F. Carr, Fla., 1934, p. 22). 

Another great chorus came Aug. 16 and 17 after a half-day's ram when 
about 2 inches of rain fell, maxima 82-91, minima 65-74. Spadefoots and 
Rana capito, subterranean species, came out to breed and Hyla squirella as 
well. 

In general this species calls even by day in rain or before an imminent rain 
(July 9) . After a downpour of a warm rain 1-5 inches they became very active. 
As every observer has remarked, heavy warm rains bring them out in mid- 
summer. Our latest record of calling came September 17. 

R. F. Deckert (Fla., 1915, p. 3) at Jacksonville, Fla., described its cry as fol- 
lows: "The cry is rather coarse, sounding like 'era, era, era/ etc. with a second's 
interval between each note." The same author (Fla., 1921, p. 22) in Dade 
County, Florida, wrote, "Their rasping calls were heard in May and June 
at igth Street, Miami, at Donn's Nursery and at 22nd Street in company. . . ." 

Breeding: This species breeds from April to August. The eggs, single on 
the bottom of shallow pools, are brown above and cream below, with outline 
distinct and jelly firm. The egg is %o-%s> inch (0.8-1.0 mm.), the inner en- 
velope ^o-Vis inch (1.2-1.6 mm.), the outer %o-M2 inch (1.4-2.0 mm.). 
The egg complement is 950. The citrine drab tadpole is small, i/4 inches 
(32 mm.), its tail long, the tail tip acuminate with a flagellum. The tooth 
ridges are %. After 40 to 50 days, the tadpoles transform from June to Sep- 
tember at %G inch (11-13 mm.). 



344 HANDBOOK OF FROGS AND TOADS 

"I have records from April 2 to August 20. Full choruses come with July 
electric storms. The males usually sing in water from half an inch to an inch 
deep at the edges of temporary pools or ditches or of temporarily inundated 
portions of pond margins. The choruses are large. Wright (1933) says that 
the call is not very loud; it has little carrying power, but at close range a large 
chorus assaults the ear-drums mercilessly" (A. F. Carr, Jr., Fla., 1940!), p. 61). 

Journal notes: We found these frogs on porches, in chinaberry trees, oaks, 
and other trees, as well as in vines around the houses, in fields and gardens 
around buildings, in open ponds, in pine barrens, in pine and oak groves, 
along roads, and in shallow roadside and pine barren pools. On June 22, 1922, 
a boy at Camp Pinckney, Ga., brought me a Scraper taken in his own house. 

July 3, 1922. Along the Folkston road, Ga., in temporary pools and ditches 
with Btifo terrestris were plenty of Hyla squirella. There is more vibration in 
the call of Hyla jemoralis. I could not hear the Hyla squirella a few rods 
away. The calls of Hyla femoralis and Bttfo quercicus drown it out. Hyla 
squirella does sometimes croak from the water surface when sprawled on the 
water. 

Okefinokee Swamp, Ga. On July 3, 1922, about 11:55 P.M., we were at 
Anna's pond. We heard the Hyla squirella at a distance between two distant 
houses. In the saw palmettos and the grass stools were many scrapers. We 
found one pair in the grass near the edge. Others were found in saw palmettos 
about the border of the pool. The same night, near Trader's Hill, we came 
to a grassy overflow pond. In a clump of bushes and saw palmetto were sev- 
eral Hyla squirella. In grassy stools in shallow water were others. 

On August n, 1922, at Camp Pinckney, Ga., 2-3 P.M., we heard several. 
At 8:30 we returned to Camp Pinckney. There were countless Hyla squirella 
males on the ground in a road filled with temporary pools, in water 1-3 inches 
deep. The vocal sac is hyaline, more or less inflated for some time. The call 
is not so fast as that of Hyla jemoralis, but swift nevertheless. Those in water 
were greenish, those at the edges of the pool, brownish. . . . We could find 
no females. Some males, though quite small, were croaking. 

Authorities 1 corner: 

W. Bartram, Gen., 1791, p. 278. A. F. Carr, Jr., Fla., 1940^ p. 61. 

P. H. Pope, Tex., 1919, pp. 96-97. G. L. Orton, La., 1947, pp. 369-375. 

A. H. Wright, Ga., 1932, p. 312. 

Common Tree Toad, Common Tree Frog, Tree Toad, Northern Tree Toad, 
Changeable Tree Toad, Chameleon Hyla, Varying Tree Toad 

Hyla versicolor versicolor (Le Qonte). Plate LXXIII; Map 20. 

Range: Maine, southern Canada, west to Manitoba, Minnesota, South Da- 
kota, south through Kansas, Oklahoma, to the Gulf States and northern Flor- 
ida (Texas and Arkansas in part only). 



TREE FROGS: HYLIDAE 



345 




LXX/7. #y/0 squirella (XO- 
Male croaking. 2,3,5. Males. 4. Female. 



LXX//7. Hy/0 versicolor versicolor. 
1,2,6. Males (X%)- 3- Female (X%)- 4- 
Males croaking (X%). 5- Eggs (X%). 



346 HANDBOOK OF FROGS AND TOADS 

Habitat: Trees, mossy or lichen-covered stone fences, decaying fruit trees. 

Size: Adults, i J /4-2% inches. Males, 32-51 mm. Females, 33-60 mm. 

General appearance: This frog varies in color from pale brown to ashy gray, 
to green; it has a granular skin, dark irregular star on upper part of the back, 
a black bar on the upper eyelid, black-bordered green bars on the legs, a black- 
bordered light spot below each eye. It might easily be taken for a stone or bit 
of bark with lichen on it. The rear of the femur has dark reticulations on 
orange. The groin, axilla, and under parts of hind limbs are orange. 

Color: Male. Belly and lower breast white or cartridge buff. Breast between 
fore limbs, which is the vocal sac, pale flesh color. Middle of throat dull citrine 
or ecru-olive with black spots. Chin like belly with blackish spots. Underside 
fore limbs like belly. Underside of hind legs and rear of thigh bright orange 
or orange-rufous. The spots on the back, one of which is an irregular star on 
upper back, are citrine-drab or deep olive with black borders. 

Female. Raleigh, N.C., from A. P. Chippey, May 12, 1950. Under parts in- 
cluding throat white becoming almost hyaline on underside of femur. Front 
of femur, groin, inner side of tibia, tarsus and first toe, and axilla cadmium 
yellow, deep chrome, or in places mars yellow. Rear of femur raw sienna, with 
numerous spots of cadmium yellow or light cadmium. Back of forelegs and 
hind legs and sides pale olive gray, pearl gray, or light mineral gray, becoming 
on back tea green or grayish olive or deep olive-buff. Star or spot in middle of 
upper back is vetiver green or grayish olive with dark grayish olive border. 
Two bars on femur, tibia, and forearm olive gray to mineral gray with dark 
borders. Oblique bar across eyelid of each eye but not meeting on meson is 
same color as star spot of back. Tympanum drab or cinnamon-drab. Spot be- 
low eye not so prominent as in male described, is olive-buff. Iris black with 
ecru-drab or light vinaceous-fawn spots. Iris rim avellaneous or vinaceous 
fawn; iris rim broken in front, below, and somewhat behind. 

Structure: Skin rough, warty; conspicuous disks on fingers and toes. 

Voice: The call is a loud, resonant trill, ending abruptly, 10 or n calls in 
half a minute. About the middle of May, at Ithaca, they are in the chorus stage. 
In the evening, all over the University hill and the hills near by, along the 
wooded ravines, in the thickety edges and woods of our marshes, we may 
stumble upon the noisy tiee toads slowly approaching the nearest breeding 
place. In one instance their resort is a pond at the end of a long hedge. Here, 
at the breeding season, every evening and sometimes after a thundershower 
by day, the males can be heard all along its length, slowly bound for the one 
objective, the pool, where some have already arrived. 

Breeding: These tree toads breed from the end of April to Aug. n. The 
brown and cream or yellow eggs are laid in small scattered masses or packet; 
of not more than 30 to 40 eggs on the surface of quiet pools, the packets loosel] 
attached to vegetation. The egg is ^>5-%o inch (1.1-1.2 mm.), the outer en 
velope indistinct, %-H inch (4-8 mm.), merging in the jelly mass, the inne 



TREE FROGS: HYLIDAE 347 

envelope Me-M.2 inch (1.4-2.0 mm.). The eggs hatch in 4 to 5 days. The tad- 
pole is medium up to 2 inches (50 mm.), tail long, scarlet or orange vermilion 
with black blotches around the edges of the crests, and with a long tip. The 
tooth ridges are %. After 45 to 65 days the tadpoles transform from June 27 
to August at l /2-% inch (13-20 mm.). 

Journal notes: June 19, 1907, Ithaca, N.Y. Ever since Sunday, June 16, the 
tree toads have been in chorus. Every day this week we have heard occasional 
tree toads and at night they were deafening when near their ponds. After 
each thundershower they liven up perceptibly. This evening at 8:45 P.M., I 
reached the Veterinary College pond. Frogs were in chorus. In 15 minutes I 
had captured 20 individuals (including a mated pair). Found them also in 
grass near by, migrating to the pond, one in the road just west. Toads were 
singing here also. Went out to cross roads. Here they were just as common. 
The log in southwest corner had tour perched on it. To show how tame and 
how dazed they are by the light, I stroked a croaking male with the lighted 
end of my flashlight 91 times without his stirring. He croaked just the same. 
I could have repeated the operation. In a tree on the north edge of the pond 
were four males, two on one limb facing each other. 

June 2i, Cross Roads Pond, n A.M. All around the pond were tree toad eggs. 
I staked out about six or seven areas of them. The packets or single emissions 
of eggs maybe 6-12 inches apart or only an inch or less. Sometimes from 
groups of eggs at more or less definite intervals one can determine the path 
of the pair. The eggs are almost invariably at the surface. Those beneath I be- 
lieve were laid before some of our rainstorms raised the water in the pond. 
They may be attached to grass leaves, plant leaves, or sometimes algae. The 
packets are from 15-35 eggs. Some of the first tree toad eggs laid here are 
now hatching. The wood frog tadpoles are beginning to develop good-sized 
legs. 

June i 3, Cross Roads Pond. Today at 1 130 found several fresh packets of 
Hyla versicolor eggs, attached to grass at edge or to Potamogeton leaves in 
mid-pond. Four to ten eggs in a packet. They were all at the surface. Water 
surface 74, water bottom 72. 

Authorities 9 corner: 

G. H. Loskicl, Gen., 1794, p. 891. G. M. Allen, N.H., 1899, p. 72. 

M. H. Hmckley, Mass., 1880, pp. 104- F. Overton, N.Y., 1914, p. 32. 

107. H. Sweetman, Mass., 1944, pp. 499-501, 

J. C. Gcikie, Gen., i882(?), pp. 217-218. R. L. Hoffman, Va., 1946, pp. 141-142. 

Cope's Tree Frog, Western Tree Frog, Chameleon Tree Frog 

Hyla versicolor chrysoscelis (Cope). Plate LXXIV; Map 20. 

Range: Southern Arkansas to east-central and southern Texas. 

'The two forms of Hyla versicolor, the pustulous (typical versicolor) and 



34 8 



HANDBOOK OF FROGS AND TOADS 




Plate LXXIV. Hyla vcrsicolor chrysosce- 
lis. i. Female (XO- 2,3. Males (Xi%). 



Plate LXXV. Hyla wrightorum (X%). 
1,3. Male with throat partly inflated 2. Fe- 
male in water. 4. Male in water. 



TREE FROGS: HYLIDAE 



349 



the smooth (variety chrysoscelis) are distributed over most of eastern Texas, 
but the present form seems to be confined principally to the extreme north- 
eastern counties and the Trinity and Brazos valleys" (J. K. Strecker, Jr., and 
W. J. Williams, Tex., 1928, p. 11). 

Habitat: Wooded stretches along creeks and rivers. 

"In Bowie County [Texas], under slabs of wood lying along the margins 
of sloughs, Mr. Williams found three specimens of chrysoscelis" (J. K. 
Strecker, Jr., and W. J. Williams, Tex., 1928, p. u). 

Size: Adults, i%-i% inches. Males, 36-43 mm. Females, 35-49 mm. 

General appearance: These frogs are a smooth-skinned version of our com- 
mon tree toad, Hyla v. verstcolor. Of a pair in hand, each has a light spot 
below the eye, legs barred, and the irregular cross on the back. The male at 
present is a light grayish tan with the pattern in green outlined with black. 
The female is gray with the pattern in dark olive and black. Both have the 
characteristic orange on the groin and concealed portions of the legs. In these 
two there is no black reticulation on the orange rear of the femur. Often the 
rear of the femur is marked with very fine spots. The dark mottling on the 
sides is very distinct. The pattern of the back is rather less massive than in the 
average Hyla verstcolor. The male seems a little smaller than the average 
Hyla verstcolor male. Sometimes the backs are green. 

Color: Male. Becville, Tex., March 27, 1925. Orange or cadmium orange on 
side of groin and front of thigh, merging into mars yellow or xanthine orange 
on rear of thighs and on the tibia. Spots on back, calla green. Background 
kildare green or mignonette green. 

Another male. Mud Creek, 9 miles north of San Antonio, Tex., taken by 
R. D. Quillm and A. H. Wright, June 24, 1930. Dorsum along sides pale olive- 
gray or pale smoke gray becoming on back olive-buff or seafoam green near 
edges of large dorsal spot. Dorsal spot in center half brown becoming near 
edge dark greenish olive or dark ivy green. One forward lateral wing of spot 
dark ivy green. Another oblique wing from rear of spot to each groin, dark 
ivy green. Rear and front of femur zanthme orange. As front of femur 
joins body color it becomes cadmium yellow or light cadmium, and this ex- 
tends along groin. Axil with touch of this color or lemon yellow. Rear of 
femur spotted with finely sized lemon yellow spots. A few or a line of these 
along rear tibial edge. Area around vent and along rear of femur a short dis- 
tance silvery white and black. Two bars like dorsal spot across antebrachium, 
one across base of third and fourth fingers, one faint one across brachium. 
Two such prominent spots on femur, two on tibia, one on tarsus, and four or 
five on foot. In front of vent from first femur bar to other first femur bar sea- 
foam green. Underside of fcmui, hind foot, some of belly russet-vinaceous or 
light russet-vinaceous. Black bar from nostril to eye. Dark area from eye 
through tympanum. Then an imperfect row of large dark spots to groin with 
smaller spots below it. Spot below eye white, ivory yellow, or seafoam green. 



35 o HANDBOOK OF FROGS AND TOADS 

Iris rim deep seafoam green; iris mainly spotted vmaceous-russet with some 
chocolate or burnt umber reticulations. 

Female. Same locality. Dorsum smoke gray. Dorsal spots and wings cepha- 
lic and caudal separate, each ivy green or lincoln green. Black stripe from 
snout to eye. Black villa from eye around tympanum halfway to groin. Below 
this a light drab band from tympanum almost to groin. Black line above in 
groin breaks into black spots with light cadmium below it. Spot below eye 
white. Other colors more or less as in male. Throat, belly, and underside of 
fore limbs white. 

Structure: Cope's original description (Hyla jemoralts chrysoscehs) is as 
follows: "Hyla femorahs Daudin. A specimen larger than the largest indi- 
viduals I have previously seen; differs also in the greater extent of the palma- 
tion of the fingers, and in the coloration of the concealed surface of the femur. 
In eastern specimens the posterior face of the femur is brown, with rather 
small yellow spots; in this form it is yellow with a blackish coarse reticulation, 
which only extends to the lower surface of the proximal half of the thigh. The 
sides have a double row of small black spots, which enclose a yellow band. 
This is probably a subspecies and may be distinguished by the name of 
chrysoscehs. One specimen as large as a large Hyla verstcolor was taken by 
Mr. Boll near Dallas" (E. D. Cope, Tex., 1880, p. 29). 

"Five specimens from Helotes were in the Marnock collection. These speci- 
mens are perfectly smooth above and resemble in every particular typical 
specimens of chrysoscehs from the type locality. This is the common Hyla of 
middle and northern Texas" (J. K. Streckcr, Jr., Tex., 1922, p. 15). 

Voice: It is a loud resonant trill. 

"This species is fairly abundant near Houston and I have the following 
notes on it: February 13, 1918. Have heard several in the past few days calling 
from trees in camp, but have not seen any yet. April 15. Collected one speci- 
men, a male that was calling from the branch of a pine tree. April 24. Found 
one calling in rain pool where H. sqturella was breeding. I heard them fre- 
quently on warm evenings, answering each other from trees in the woods 
near camp. After the first of May they became silent and I heard and saw no 
more of them for the season. If they laid eggs in any of the pools near camp 
they made no such noise about it as they do in the North" (P. H. Pope, Tex., 

i9'9> P- 95)- 

"2/17- Heard Hyla v. chrysoscehs calling tonight evening, ist this spring. 

"Feb. 18. Cold. No frogs or toads. 

"Specimen 1948. H. v. chrysoscehs. 7/15/44 from damp ground under wa- 
ter spigot at house 3" deep. 

"Somerset, Tex. y 1 /:* mi. S.W. 1944. Sept. 2. 

"Specimen 1951-1956 inc. Hyla v. chrysoscehs. 

"I have been hearing a strange frog or (toad) note since Aug. 27, from the 
Geo. Foster tank or pond about % mi. S.E. of our house. There had been 



TREE FROGS: HYLIDAE 351 

much rain from Aug. 22 to Aug. 28, clear since 8/28. Tonight I went to the 
tank and heard a few of these calls. After searching I found them to come 
from trees and bushes and a tree fallen in the water. I caught two that I 
thought were the croakers. Hylas, somewhat small. Finally I was watching 
two Hylas, one was croaking the other about 6" away. Female was silent, sud- 
denly the croaking male began calling the strange notes: uttered fully twice as 
quick as the ordinary notes and an altogether different tone, clear, shriller, 
2 to 4 notes; ('qui qui qui') sometimes there was a low gurgling preceding 
the call notes; most times the notes were low but now and then they could 
be heard a long distance, clearly at our house, % mile away. This satisfied me 
that the strange notes were made by the male H. v. chrysoscelis during mating 
times. All the males (6) heard that I saw were near females. On a horizontal 
limb ($" diameter) 6 feet up were 5 hylas, 2 females, } males. They were close 
together, all five within 10 inches. One male uttered these calls. It was along- 
side of a female, 'nudged* the female a few times. This pair collected, as was 
the pair first watched and 2 other single females (?) caught near where these 
calls heard. I have heard these strange notes a few times before" (notes, A. J. 
Kirn, Somerset, Tex., 1944). 

Breeding: They breed from middle March to July. The tadpoles transform 
at % inch (16 mm.). 

"This spring I found this tree-toad breeding in small rock-bound pools in 
a gravel pit. The tadpoles were light yellow. Specimens collected April 21 st 
had the hindhmbs well developed" (J. K. Strecker, Jr., Tex., 19103, p. 118). 

Journal notes: March 24, 1925, Beevillc, Tex. The air is resounding with 
Psetidacris. We heard a few scattering Hyla versicolor along the roadside 
ditches or in the distance and captured three males, but later lost one. They 
are along these roadside ditches but not in them. They look quite green at 
night. We captured one on the bole of a mesquite near the pond. Meadow 
frogs are croaking loudly in these ditches and narrow-mouthed toads arc 
common. 

June 14, 1950, Becville, Tex., at night. We were about to leave the pond 
when we heard a H. versicolor. We started for it when we heard two more 
at a distance. One we caught. It has spotting on the rear of the femur, very 
fine spots down the outer half of the femur. When we reached the house we 
lost the H. versicolor. 

June 22. We went out with R. D. Quillin. At Mud Creek, 9 miles north of 
San Antonio, we heard many Microhyla olivacea, Rana piptens, a few Bufo 
valliceps, and several Hyla versicolor chrysoscelis. We sought the tree toads. 
The first one was too high in the oaks. The next one we picked up on mud 
near the water. Then we heard a croaker near the spot but the one on the mud 
proved a ripe female. In another oak tree we found a male, 3 feet up. This we 
put with the female. When we returned to the house they were mated axillary 
fashion. 



352 HANDBOOK OF FROGS AND TOADS 

June 26, Waco, Tex. I saw some of Strecker's H. v. chrysoscelis. He pro- 
nounced my San Antonio live Hyla versicolor, H. v. chrysoscelis. These are 
smooth and quite greenish. In three H. v. chrysoscelis in the Baylor collection 
the suborbital spot is obscure or absent or present on one side and absent on 
the other. The femoral reticulation is more pronounced than in my H. v. 
chrysoscelis from San Antonio. 

Authorities' corner: 
J. K. Strecker, Jr., Tex., 19103, pp. 117- 

118. 

"In the southern and western specimens there is a tendency to the replacing 
of the brown reticulation on the yellow ground of the posterior face of the 
thighs by a number ot subcircular golden spots in the brown ground, as in 
the Hyla jcmoralts, although northern specimens sometimes show traces of 
it. This is very evident in specimens from Prairie Mer Rouge and Tangipahoa 
River, Louisiana, and Dallas, Texas. As a general rule, too, the portions of 
the limbs concealed or in contact with each other when flexed, are in northern 
specimens more fully marbled with yellow, even covering the whole inner 
surface of the tibia and the light interspaces more or less angular, while in the 
Stntlisca bandmii and the southern and western specimens of H. versicolor 
the amount of marbling is less, and the interspaces are often reduced to small 
circular spots. I have, however, been unable to characterize them as more than 
a variety, to which I give the name of H. v. chrysoscelis'' (E. D. Cope, Gen., 
1889, pp. 374-375). 

"It is easy to be confused in trying to solve Cope's original Hyla versicolor 
chrysoscelis, H. v. versicolor, and Hvla v. phacocrypta. Viosca positively as- 
serts that H. v. phaeocrypta type is H. v. versicolor. Stejneger early pointed 
out that H. v. chrvsoscelis could not be H. fcmoralis. 

"When after H. avivoca with Viosca and Chase on June n, 1930 in Tick- 
faw and Tangipahoa Country, 'we took in road ditches several H. v. versi- 
color. Are they H. v. chrysoscelis? Quite smooth. The female is very large in 
contrast to the males captured. Started for H. avivoca nearby. It is not the 
H. versicolor of Louisiana, but is quite different and different in ecological 
habits. Yet if I read Cope's recharacterization (1889) of H. v. chrysoscelis 
(1880) (partially from Louisiana, Tangipahoa River this very country I am 
in) one might suspect Cope might have had some H. avivoca. If so his Tangi- 
pahoa River, La., and Dallas, Texas, description is of a composite form. Cope 
(1880) put his form in H. femoralis, but Hyla avivoca never was H. femoralis 
in habit. It is found in tupelo gums, H. femoralis in pine woods. Each of these 
and H. versicolor are in this region of Louisiana. . . .' Strecker redescribes 
H. v. chrysoscelis. He calls it a smooth-skinned frog with a yellow suborbital 
spot. He also alludes to Cope's character of fine femoral spots. I have taken 
H. versicolor from Beeville to north of San Antonio. I believe that it is the 
form Strecker has interpreted as H. v. chrysoscelis. I showed him some live 



TREE FROGS: HYLIDAE 



353 



specimens from Mud Creek 9 miles north of San Antonio, Texas. These 
seemed much like Waco, Texas, specimens he showed me. Both lots (mine 
and his) are smooth-skinned but some have no suborbital spot, or occasionally 
they have a line from nostril to eye and backward. The Louisiana material 
needs to be restudied. At present we will hold to the Dallas (Boll) material as 
the center of its (H. versicolor chrysoscehs) eastern Texas to Arkansas range" 
(A. H.Wright, MS, 1930). 

Recently H. M. Smith and Bryce C. Brown assigned the Hyla versicolor of 
the "Lower Mississippi Valley west through eastern Texas to about Leon and 
Austin counties" to H. versicolor chrysoscelis. This they diagnose as follows: 
"Like H. v. versicolor except rear surface of thigh more extensively dark 
marked, leaving only isolated, circular light areas of moderate size." This 
seemingly is in agreement with Cope's saying he had it from Dallas, Tex., 
and Tangipahoa River, La. The new form, H. versicolor sandersi, after our 
friends Mr. and Mrs. Ottys Sanders, they restrict as follows: "The Balcones 
escarpment and its vicinity in central Texas, south at least to Atascosa County, 
east to Bastrop County, north to Travis County and no doubt to McLellan 
County." "The Texan subspecies may be diagnosed as follows : rear of thigh 
light (orange in life), with fine white flecks, entirely lacking dark marks ex- 
cept at extreme medial border; fingers not or only barely perceptibly webbed; 
skin smooth; maximum snout-vent length lesser, 4? mm. in males, 48 mm. in 
females. . . ." 

Dusky Tree Toad 

Hyla versicolor phaeocrypta (Cope). Plate LXXVI. 

Range: Specimens from Nashville, Tenn., Olive Branch, 111., Mt. Carmel, 
111., Olney, 111., Gull Lake, Brainerd, Minn., and Springfield, S.D., have been 
assigned to this form. 

Habitat: River valley and lake shore. 

Size: Adults, i%-i% inches. Males, as large as 33 mm. Females, as large 
as 36 mm. 

General appearance: This is another of Cope's H. versicolor-H. femoralis 
puzzles based on preserved material. It is as yet unsolved; however, it is 
probably not a good form. Cope's original description follows : 

"A single specimen of a strongly marked variety of this species was sent 
to the National Museum from Mt. Carmel, 111., by Lucien M. Turner (No. 
12074). It is smaller, having the average dimensions of H. femoralis. The color 
is a dark brown, with three rows of large approximated darker brown spots. 
The groin and concealed faces of the thigh are yellowish brown, with a very 
scanty speckling of darker brown, very different from the usual coarse netted 
pattern. At first sight one suspects this to be a specimen of Hyla femoralis, 
but it possesses all the essential characters of the integument and feet of the 




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TREE FROGS: HYLIDAE 355 

H. versicolor, as pointed out in the analytical table of the genus, including 
also the light spot under the eye. It may be called H. v. phaeocrypta 9 (Cope, 
Gen., 1889, p. 375). 

In hurried visits to three of its localities, we have met questionable forms. 
Live material from Wabash Valley is our greatest need. Viosca now pro- 
nounces the type of Hyla phaeocrypta a poorly preserved Hyla versicolor. 
What is this form? In the last fifteen years little has appeared to add to the 
solution. 

From Itasca Park, Minn. This is a smooth-skinned tree toad, gray with 
prominent crossbars on legs and varying types of spots on the back. In some 
these spots assume a cruciform pattern or are arranged in three or four rows. 
There is a white or ohvine spot below the eye. From the eye backward ex- 
tends an oblique band of gray or vmaceous bordered above with black. There 
is much cadmium orange or cadmium yellow on rear and front of femur, on 
underside of foot, in groin, and in axil. From Walhalla, N.D. In darkness 
and moisture, this frog was dark grayish olive and very little dorsal pattern 
showed. Brought into the light, he is rapidly becoming lighter and the dorsal 
pattern is coming out. He lacks the cruciform patch of typical versicolor. 
There are about four rows of irregular spots on the back. 

These and the Kansas, South Dakota, and Wisconsin records are likely to 
prove variant H. v. versicolor. 

Structure: For those who wish to follow the relative measurements of Hyla 
avtvoca, Hyla v. versicolor, and Hyla v. phaeocrypta (Illinois northwest- 
ward), the following table ot 1930 is given: 

H.a. H.p. H.v. H.a. H.p. H.v. 

No. USNM 75026 70644 C.U. 75018 70643 3636 

Sex Male Female' 5 Female Female 

Length 28 28 28 36 36 36 

Head-tympanum 10 9 10 n 12 12.5 

Head-angle of mouth 9.5 7 9 10 10 11.5 

Width head n 10 12 12 13.5 14 

Snout 5 4.5 5-5 5-5 6 6 

Eye 334 4-5 3-5 4 

Interorbital space 3.5 3.5 2.5 5 4 5 

Upper eyelid 3.5 2.5 2.5 4 3.5 4 

Tympanum 2233 2.5 2.5 

Intertympanic space 10.5 8.5 9 n 11.5 11.5 

Internasal space 3.5 333-53 4 

Fore limb 18 14 20 20 17 24 

ist finger 4.5 4 3-5 5-5 4-5 6 

2nd finger 6 4.5 5 7.5 5 8 -5 

3rd finger 9.5 6 8 10 7 n 

4th finger 7.5 5 6 8.5 6 9 

Hind limb 42 35 44 55 53 54 



356 HANDBOOK OF FROGS AND TOADS 

H.a. H.p. H.v. H.a. H.p. H.v. 

Tibia 15 15 16 18.5 19 

Foot with tarsus 19.5 16 24.5 24 27 

Foot without tarsus 14 9 14 14.5 14 15.5 

isttoe 4-5435 6 7 

2nd toe 7 6 4.5 8 8 10 

3rd toe 10.5 8 8 12.5 10 12 

4th toe 14 9 ii 14.5 12 15.5 

5th toe ii 8 8 12 9.5 12 

Remarks: The material which comes into this consideration is: 
USNM no. 12074, acl - Mt - Carmel, 111. L. M. Turner (type). 
USNM no. 70642, 34 mm. 3 Gull Lake, 10 mi. n. Brainerd, Minn. 
USNM no. 70643, 36 mm. $ Gull Lake, 10 mi. n. Brainerd, F. H. Uhler. 
USNM no. 70644, 28 mm. yg. Gull Lake, 10 mi. n. Brainerd, F. H. Uhler. 
USNM no, 71588, 10 mi. ne. Olney, Richland, 111., Robt. Ridgway. 
USNM no. 68719, 28 mm. yg. or ? Springfield, S.D., E. C. O'Roke. 

Mittleman kindly showed us his manuscript. It is a good paper and he may 
be right. In the preceding table we compared one 28-mm. and one 36-mm. 
specimen of each form, six examples in all. We still retain H. v. phaeocrypta 
in this book for someone to catch the Wabash frog in the act and then for the 
same person the same season to familiarize himself with the song of H. avi- 
voca. 

Some of Mittleman's statements and our comments follow: 

1. "In addition I have heard the call of this species (H. phaeocrypta, i.e., 
avivoca) in the gum trees of the mud flats along the Wabash River on the 
outskirts of Terre Haute, Indiana" (p. 34). Quite likely he did hear it. Why 
compare this memory with an old preserved specimen ? 

2. "The highly distinctive voice of phaeocrypta (avivoca) need be confused 
with no other North Ameiican hylid save possibly H. cruet fer" (p. 35), Too 
comprehensive a statement. Authors speak of "breeding song," "rain song," 
"chorus note," and other notes for one species. One or two species may vary 
an octave, 

3. "Phaeocrypta. Call, a piping, bird whistle. Versicolor. Call, a toad-like 
trill" (p. 36). To DeKay versicolor had a ventnloqual quality, to Cope a reso 
nant trill, to Fowler a jerking cry, to Dickerson a cat purr or lamb bleat, tc 
Overton a musical trill like a low-pitched whistle (trill and whistle combined) 
and a turkey call, to Harper (1921) a vigorous high-pitched trill, and so on 
Did Turner actually catch a birdlike Hyla at Mt. Carmel? What is birdlike 
Overton's turkey root, Viosca's and Wright's woodpecker call for avivoca, o\ 
somebody's musical thrushlike notes ? What birds do we choose ? 

4. "If Harper's specimen is truly referable to phaeocrypta it probably indi 
cates a significant trend in Atlantal coastal populations. It seems much mor< 
likely that it is simply a versicolor" (p. 34). Harper at Louisville, Ga., took , 



TREE FROGS: HYLIDAE 357 

female avivoca of 49 mm., 5 mm. longer than what we reported in 1933, or 
6 or 7 mm. more than Mittleman's specimens. Why not? We have been in 
the field many times with the four veterans who know H. avivoca best, 
P. Viosca, Jr., O. C. Van Hyning, A. F. Carr, Jr., and F. Harper. They know 
the southeast frogs alive, in alcohol, and theoretically. 

Color: Male. Itasca Park, Minn., Aug. 26, 1950. The spots of back are deep 
olive-gray with dark margins. These spots consist of four irregular dorsal 
rows, and a bar from each eyelid extending obliquely backward and inward 
but not meeting its fellow. The background of the back is smoke gray with 
black line from snout to eye, resumed back of eye, and extending over tym- 
panum and arm insertion and along side. The area below it and involving the 
tympanum is drab-gray or light purplish gray. This area on its lower portion 
is separated from the light cadmium of the axil by oblique black border. The 
spot below the eye is white. Transverse area above vent is white, bordered 
below by black. Below this is a pale flesh area. Rear of femur has prominent 
cadmium orange spots with amber brown interspaces. Underside of tibia is 
amber brown with a few orange spots. So also is the inside of tibia and foot. 
The groin is mainly cadmium yellow. The eye is spotted with congo pink. 
The throat is light grayish olive with rest of venter white. 

Another male. Walhalla, N.D. Sept. i, 1930. The general dorsal color is dark 
olive-gray, becoming olivine on top of foreleg, on top of femur and tibia, and 
on sides above the dark edge of lateral band. The spot below the eye is of the 
same color. Band from eve through tympanum and over arm insertion is light 
drab. The tympanum is grayish olive. This lateral band is bordered by black. 
Just above groin is a prominent black-edged spot; its center is olivine. There 
is a slight touch of clear orange or cadmium yellow in axil, groin, and front of 
femur. The rear of femur is finely spotted with cadmium yellow, on a xan- 
thme orange or amber brown background; same on underside of tibia and 
inner side of foot. The femur is slightly barred, the tibia more so. There is a 
V from one eyelid to the other. The spots of back are black or with citrine cen- 
ters. The underside of lower jaw is olivine with some dark clashes. The lower 
throat is deep grayish olive. The rest of under parts are white except for im- 
dersurfaces of hind limb, which arc pale flesh color. Just above anus is a light 
lumiere green area bordered below by black, and this in turn by pale grayish 
vinaceous, below that a silvery pebbling, some of which appears also on tarsus. 
The iris is black, heavily spotted with vinaccous-tawny; pupil rim broken 
above, below, behind, and in front; pupil rim is light greenish yellow above. 

Female. Itasca Park, Aug. 26, 1930. The spots on back are light olive-gray or 
tea green outlined with dotted lines of black. The bars on femur, tibia, and 
antebrachium are the same color. The general background of back is pale 
drab-gray. From the eye there extends backward along the side, an ecru-drab 
or pale brownish drab band bordered above by black-dotted line. The spot 
below the eye is white or marguerite yellow. The tympanum is light cinna- 



358 HANDBOOK OF FROGS AND TOADS 

mon-drab. The front of femur, underside of tibia, and rear lower half of femur 
is cadmium yellow with same color in groin and to slight extent in axil. This 
cadmium yellow forms fine spots on rear of femur and slightly on rear and 
forward edge of tibia, with interstices of sudan brown or antique brown. The 
iris is heavily spotted with japan rose. Ventral surfaces are white. With this 
frog, the spots on the back were a large irregularly cruciform one on the for- 
ward part of the back and two smaller ones to the rear and laterad. 

(These three specimens, two from Itasca Park, Minn., and one from Wal- 
halla, N.D., may eventually, of course, be referred to H. v. versicolor.) 

Voice: "As it is impossible to revive the voice of Copy's type specimen, some 
one must hear and collect its descendants in or near the type locality. Fortu- 
nately, this has virtually been done through the kindness of Mr. Karl P. 
Schmidt, of the Field Museum, who has called my attention to a series of 
specimens from Olive Branch, Illinois, in that museum, and these' seem to be 
the same as Cope's phaeocrypta. The field notes of Mr. C. M. Barber which 
were submitted by Mr. Schmidt are especially interesting, as they may supply 
the missing evidence: 

" Whilst collecting Hyla cincrea with a light in a boat, on Horseshoe Lake, 
May 23, 1907 with Dr. Chatham, my attention was called to a bird-like cry 
coming from trees and bushes overhead. At first I thought it resembled the 
rattle of Acns, but it has a bird-like quality quite peculiar. Several searches 
were unsuccessful, but finally we were able to find the sober-colored little 
frog when he uttered his cry, fearlessly, within a few feet, and with the light 
fully on him. Their note is strong and penetrating and more pleasing than 
any Hyla that I know. I had previously taken one in the grass near the lake, 
but mistook it for a juvenile versicolor " (P. Viosca, Jr., La., 1923^ pp. 97-98). 

In 1924 Robert Ridgway (111., p. 39) writing under the title, "Additional 
Notes on Hyla phaeocrypta ( D )," reminisced as follows: 

"Mr. Viosca's observations on Hyla phaeocrypta Cope in Copeia 122 recall 
to my memory an experience o many years ago in the bottom-lands of the 
Wabash and tributary streams near Mt. Carmel, 111. The bird-like notes de- 
scribed by Mr. Viosca were frequently heard by me in the woods, so distinctly 
bird-like that I was constantly looking for the unknown bird; in fact, there 
was a "standing reward" offered to my boy friends for a specimen never dis- 
covered. Some years later, when my father moved to a farm near Wheatland, 
Ind., I learned from him that it was a tree-frog. Whether Hyla phaeocrypta 
or not, the species is so abundant in the bottom-land woods that at certain 
seasons (if my memory is not at fault, in late summer or autumn) its clear, 
loud piping notes as different as possible from the croak of H. versicolor 
were the dominant and often the only sound to be heard" (R. Ridgway, 111., 
1924, p. 39). 

Breeding: Nothing of record. 

Journal notes: Sept. 13, 1929, Calhoun, 111. While collecting in the northern 



TREE FROGS: HYLlDAE 



359 



gopher frog areas which Lee Ackert showed us, we came to a pond. When 
about to leave, we heard a Hyla versicolor. It was in a bush in the deep water 
of the pond, and so distinctly H. v. versicolor that we passed it by in our 
search for R. areolata circulosa. Later we wondered if it might not have been 
Ridgway's H. phaeocrypta. 

Sept. 14. Went to Mt. Carmel, 111., and New Harmony, Ind., for a day but 
heard nothing like the reputed H. v, phaeocrypta. 

Aug. 26, 1930, Gull Lake, Minn. Wanted to take some of the queer hylas 
F. M. Uhler secured here, but found instead almost every other species of the 
region. 

Sept. 7, 1930. Drove around the country at Mt. Carmel and listened at night 
and by day, but discovered no queer Hyla notes such as those of H. v. phaeo- 
crypta are reputed to be. 

Oct. 7, 1929. A small male tree toad from northern Wisconsin (from Caro- 
lyn Weber), we provisionally placed in this group. Our general notes on it are: 
This is a small specimen of tree toad. It is brownish gray on the back with 
irregular black markings, forming an irregular and imperfect cross. A dark 
band extends from eye downward across the tympanum and along the side. 
The top of this band is marked with a broken black line bordered above by 
light. The sides are speckled brown. The legs are barred, the throat is dark. 
There is orange on rear ot hind legs, in groin, and in axilla. 

Not until we have several of these frogs alive from Mt. Carmel, Olive 
Branch, and Gull Lake, will we feel satisfied or competent to weigh the posi- 
tion of this species. Is it a separate form or H. versicolor versicolor, H. v. 
chrysoscelis, H. avivoca, or Pseudacns brachyphona? 

Authorities 9 corner: In 1923 while provisionally placing his later H. avivoca 
under H. phaeocrypta, Viosca wrote: "As virtually all the distinguishing char- 
acters of my new frog were lost in the preservative, its general resemblance to 
Hyla versicolor made comparison with museum material difficult, especially 
since versicolor is such a variable species. Except for the smaller size, which 
could easily have been due to age, there is nothing in Cope's description of 
phaeocrypta that would suggest my specimens. Dr. Stejnegcr has compared 
specimens of my species with Cope's type of phaeocrypta with the result that 
structurally at least he could not tell them apart, yet this in itself was not con- 
clusive, as my specimens also exhibit a structural resemblance to small speci- 
mens of Hyla versicolor chrysoscehs. . . . Hyla phaeocrypta . . . should how- 
ever be rediscovered in the type locality and the voice noted. Although there 
are apparent difTerences in the specimens at hand, between the phaeocrypta 
of Illinois and the bird-voiced Hyla of Southeast Louisiana, it is difficult to 
pass an opinion with the limited material available. These also must be con- 
sidered the same until both forms can be compared in life, or better, be heard 
by the same person" (P. Viosca, Jr., La., igijb, pp. 97-98). 

In describing his Louisiana Hyla as Hyla avivoca not Hyla phaeocrypta 



360 HAKDfiOOK OF FROGS AND TOADS 

Viosca said in 1928: "The species described herein had been called tentatively 
Hyla phaeocrypta Cope pending its comparison by the writer with Cope's 
type of phaeocrypta, as well as with other North American Hyla types in 
the National Museum. I have had an opportunity recently to make such a 
study in the National Museum and found the Louisiana form to be a species 
distinct from any North American Hyla heretofore described. Further, I 
found the type of Hyla versicolor phaeocrypta Cope, U.S.N.M. No. 12074, to 
be a fairly typical specimen of Hyla versicolor, poorly preserved as to texture 
and color, but well within the range of individual variations normally ex- 
hibited by that species. The typical cruciform pattern of versicolor, though 
faint, is readily discernible, and the structural characters pointed out by Cope 
place it unquestionably with that species" (P. Viosca, Jr., La., 1928, p. 89). 

Note on W. T. Neill's (1948) significant paper on an eastern race, Hyla 
phaeocrypta ogechiensis for Georgia and Carolina specimens: Having visited 
the type locality of Hyla phaeocrypta without success, we want evidence from 
someone with success in Mt. Cartnel and the lower W abash country. Northern 
specimens arc doubtless versicolor but avwoca-phaeocrypta is not yet proved. 
Avivoca's range may extend in the Upper Coastal area to North Carolina. 

Sonora Tree Frog, Wrights' Tree Frog, Sonora Hyla 
Hyla wnghtontm Taylor. Plate LXXV; Map iS. 

Range: North-central Arizona, New Mexico, Chihuahua possibly to Texas. 

"To date the author has found this frog only on the southern forested edge 
of the Colorado Plateau from the vicinity of Williams east to McNary. The 
full range extends cast into Texas and south into Mexico. 

"The Colorado Plateau extends roughly east and west through central Ari- 
zona into New Mexico. The altitude is approximately 7000 feet above sea level. 
In general the southern part is in the transition zone and is covered by typical 
vegetation including a broad belt of ponderosa pine with small areas of Doug- 
las fir, white fir, Mexican white pine, and other irees. . . . No specimens of 
Hyla wrightontm were found in the isolated Final Mountains, south of Globe, 
although they reach fiom upper Sonoran through the transition zones, rang- 
ing from 4000 to 8000 feet in elevation. No frogs were found in the large 
springs at Pinetop, whose waters have a temperature of about 50 F. The 
waters of open ponds arc warmed during the day, and remain at a higher 
temperature. 

"The lower limit of the distribution of the species in this region seems to 
be defined by the rim of the Colorado Plateau, with an elevation of 5000 feet. 
The 1936 season was spent at Indian Gardens, 22 miles east of Payson, in an 
area directly beneath the rim of the plateau (here called the Mogollon or 
Tonto Run). No specimens were found in this well forested area, though 



TREE FROGS: HYLIDAE 361 

frogs were found on the plateau above on several trips. Hyla eximia was not 
listed for the Grand Canyon National Park. . . . 

"Judging from the small number of specimens found, Williams is probably 
the western edge of this frog's range. Although the summer rains were few 
in 1937, they were heavy enough to have brought out all individuals, but very 
few were found. Many suitable ponds were completely barren. Continuing 
east numerous specimens were found along the rim. At Hart Canyon, which 
is 50 miles south of Winslow, and at Pinetop and McNary the numbers in the 
summer ponds were very great" (W. L. Chapel, Ariz., 1959, p. 225). 

Habitat: "Before and after the breeding season the frogs are found occasion- 
ally throughout the forest. Clever camouflage and ventriloquism aid them in 
escaping detection. They are found on the ground in damp places and in the 
trees. Twice the author found several which had been jarred loose when a 
tree was felled. In one of these instances, a frog fell from the top of a tree 
about 75 feet high. On sultry days they were heard calling from the trees" 
(same, pp. 225-226). 

For breeding they seek large, shallow, grassy, rain-water pools, permanent 
ponds, new ponds, brooks, even wells. 

"It is a species apparently adapted to semidescrt conditions" (E. H. Taylor, 
Ariz., 1938, p. 439). 

Size: Adults, 1-1% inches. Males, 24-44 mm * Females, 24-48 mm. 

General appearance and Color: At first gl.mce, these frogs look like Ander- 
son's tree frogs (Hyla andersonii), green Pacific tree frogs (Hyla regtlla), or 
large chorus frogs (Psettdacni). They are bright green with a dark purple 
or black line from the snout through the eye, broadening over the tympanum 
and extending halfway along the sides. This vittal stripe may be broken into 
groin spots, is slightly margined above with white, on the sides is irregular in 
outline, and may be lighter and bronzy in the center. There are two conspic- 
uous linear black spots extending backward from the rump, occasionally 
paired dark spots on the forward part of the back, but no marks between the 
eyes. The groin is conspicuously greenish orange or old gold, as is the rc.ir 
of the femur, which is unspotted. There arc irregular spots or bars on the 
tops of the arms and legs. There are irregular dark borders on the upper and 
lower jaws ending in a dark spot at the shoulder. The throat of the male is 
dull greenish tan, that of the female white. 

"This species is related more closely to H. regilla and H. lajrentzi than to 
the typical H. eximia. From the former it differs in having a smooth rather 
than pustular skin, and in having a longer leg, the tibiotarsal joint reaching 
the tip of the snout or beyond, instead of to the region of the eye. The webbing 
of the toes is somewhat less and the diameter of the tympanum is greater than 
half the diameter of the eye; the toes and fingers are wider with somewhat 
wider pads. 



362 HANDBOOK OF FROGS AND TOADS 

"From H. lajrentzi it differs in having the webbing between the toes some- 
what less with narrower fingers and toes, larger choanae, a shorter, blunter 
snout, somewhat deeper in front ot nostrils. The front edge of the tibia is 
heavily-spotted with brown, instead of having it blackish with a cream-white 
or silver line which is continued to foot" (E. H. Taylor, Ariz., 1938, p. 439). 

Structure: Skin smooth above, granular below; a fold of skin across chest; 
a fold across the base of the throat in the female; fingers and toes with well- 
developed disks; the fourth toe very long; fingers slightly webbed at the base; 
toes % to l /2 webbed; tympanum half the diameter of the eye; tibia half the 
length of the body; a tarsal fold present; inner metatarsal tubercle present, 
but outer metatarsal tubercle present, inconspicuous, or absent. 

"Description of the type. A medium-sized member of the Hyla eximia 
group. The snout is rather truncate or bluntly conical, with the canthi more 
or less distinct but rounded; the line between eye and nostril somewhat con- 
cave, sloping obliquely from canthus to edge of lip; diameter of eye somewhat 
greater than distance of eye to nostril, and equal to distance of nostril to mid- 
dle of upper labial border; nostrils below edge of canthus, the distance be- 
tween them about equal to their distance from eye; the area about nostril 
slightly elevated, and a slight, shallow groove present between nostrils; diame- 
ter of the tympanum is contained in the diameter of the eye slightly more than 
1.5 times; the distance between the tympanum and eye about .65 of diameter 
of tympanum. 

"Tongue broadly cordiform or subcircular with a very slight median 
emargination posteriorly; free posteriorly for two fifths of its length. In males 
the openings ot the single vocal sac are lateral to the tongue and much elon- 
gate; tongue papillae not prominent; the raised prominences bearing the 
vomerine teeth are large, placed slightly diagonally and closer to each other 
than to choanae; they arise near anterior level of the choanae, but do not reach 
their posterior level. The openings of the mucous glands form a continuous 
groove anterior to choanae; latter proportionally large. 

"A vestige of a web between first three fingers, but practically obsolete be- 
tween outer fingers; disks on the fingers moderate, only a little wider than the 
toes, the widest one on outer fingers equal in width to a little more than half 
the diameter of tympanum; first finger reaching to a point halfway between, 
the distal subarticular tubercle and the terminal disk of the second; the distal 
subarticular tubercles large, that on outer finger very slightly bifid on right 
side (probably abnormally); a slight dermal fringe on the lateral edges of 
fingers; fourth finger longer than second. 

"Legs elongate, the limb laid forward, the tibiotarsal articulation reaches to 
the tip of the snout or beyond slightly; when limbs are folded at right angles 
to body the heels overlap about two millimeters; terminal disks on toes not 
wider than digits, distinctly smaller than finger disks; a well-defined tarsal 



TREE FROGS: HYLIDAE 363 

fold; a prominent, salient, inner metatarsal tubercle, its length in the first 
finger length about two and one-half times; outer metatarsal tubercle distinct, 
flattened, lying behind the anterior level of the inner tubercle; inner toes 
webbed at base, the depth of web from one fourth to one third the length of 
the outer toes; the web between the three outer toes incised to a point one third 
the distance between the two proximal subarticular tubercles of the fourth 
toe; supernumerary tubercles on palm and foot more or less distinct. (In males 
the large tubercle at the inner part of the base of first finger is covered with a 
corneous callosity, usually very light brown in color.) Anal flap rather wide, 
not especially modified; no axillary web; skin on body relatively smooth, un- 
der magnification one observes minute corrugations, more evident above eyes; 
a strong skin-fold across the breast; ventral surface granulate, the granules 
on the anterior part of abdomen largest, less distinct on throat and chin; gran- 
ulations prominent on median ventral, and to some extent, posterior part of 
thighs; a rather thick but relatively indistinct fold above tympanum" (E. H. 
Taylor, Ariz., 1938, pp. 437-438). 

Voice: The call is a low-pitched, harsh, metallic clack, with no trill, and 
consists of two to ten, twelve, or more notes given in succession. These may 
be speeded up toward the end of the call. The vocal vesicle is very large, single, 
and transparent, with less yellow or green in it than is found in most tree 
toads. 

"Fresh rains bring out a renewed chorus. A chorus usually lasts two or 
three nights and then quickly thins out to a few disconsolate males for two to 
four more nights. A chorus continues all night, with the greatest volume be- 
fore midnight. Probably new individuals make up most of each new chorus" 
(W. L. Chapel, Ariz., 1939, p. 226). 

Breeding: July 9, 1942. These tadpoles range from 28-40 mm. in length 
(average 32 mm.) with body lengths of 11-17 mm - (average 13.5 mm.) and 
tail lengths of 17-23 mm. (average 19 mm.). 

Tadpole. Hart Canyon, CCC Camp, Winslow, Ariz., July 16-24, J 933- Hotly 
dark olive or deep olive clotted with closely set mignonette green, light yellow- 
ish olive or dull citrine dots; the snout is almost the color of these dots with 
no olive. The deep olive of the back extends along as a rim margin on the 
upper edge of the musculature. There is a tendency for a similar rim on the 
lower edge of musculature. The middle portion of the musculature is olive 
lake, olivc-ochcr, or old gold. The hind legs as they begin to develop have 
much the same color. The throat also is this color. The sides of the belly have 
a flesh color or light vinaceous-cmnamon cast. The middle of the belly is 
white. The tail crest is finely dotted on the back with some tendency toward 
reticulation of the dots. The lower crest to the naked eye may look to be almost 
transparent but is actually covered all over with very fine deep olive dots. 
Pupil rim of eye is salmon colored, succeeded by black iris flecked with light 



364 HANDBOOK OF FROGS AND TOADS 

viridine yellow and flesh color. Two transformed at 15 and 13.5 mm. They 
were green at transformation and had a suggestion of a dark indefinite area 
or band ahead of the eye. 

"The breeding season is from June to August, but in any year the dates vary. 
The determining factor is the heavy summer rains common on the Colorado 
Plateau in July and August. These storms delimit the breeding season. The 
seasons noted were: 1933 Pinetop and Hart Canyon, 7000 feet, July 2-Aug. 9, 
heavy rains; 1935 Pinetop, 7000 feet, July 7~July 28, normal rains; 1937 Wil- 
liams, 7000 feet, July 2-July 14, subnormal rains. Breeding is intermittent 
within this period" (W. L. Chapel, Ariz., 1939, p. 226). 

Two ripe females 37.5 and 42 mm. were taken at Santa Fe, New Mexico, 
June 1 8, 1874, by H. W. Henshaw. 

"The migration of frogs toward fresh rain-water pools begins as soon as 
the summer rains start. . . . The numbers of specimens present indicate that 
large, grassy, shallow ponds are quite common in the typical open parks in 
the ponderosa pine forests in central Arizona. The grass provides resting 
places for the frogs. In open water a few may swim about, but the majority 
remain along the edge, resting on the bottom. . . . 

"The duration of the egg and tadpole stages is unknown. Some eggs were 
tagged near Pinetop, but the author was transferred the next day. No eggs 
have been found elsewhere. Some tadpoles were coralled several times, but 
subsequent freshets removed the broods. Many choruses and many tadpoles 
have been found in rapidly flowing brooks but no eggs have been found. 
Probably the egg masses either are laid in quiet back-washes or are washed 
into them to hatch. The frequent freshets cause high mortality, and after the 
rainy season as the brooks dry up many tadpoles are isolated in pools and are 
dried up with the water. 

"Frequently in small pools the water is nearly black with the swimming 
tadpoles. The tadpoles gather in very shallow, warm side-pools full of de- 
caying vegetable matter. They also gather in great numbers around fresh cow 
manure and apparently feed on the dissolved and softened material. In areas 
where this species is abundant, in later summer, when the tadpoles transform, 
the ground around the ponds is covered with out-going froglets. These vary 
in length from 10 to 13 mm. Many leave the water still carrying stubby tails" 
(W. L. Chapel, Ariz., 1959, p. 226). 

Nov. 27, 1945. Again Bill Chapel arrives in the "nick-o-time," to tell us that 
those eggs he tagged near Pinetop were in small loose masses possibly the 
size of a small teacup loosely attached to grass stems just below the surface 
of the water. 

Journal notes: On June 10 or 12, 1933, as Mr. W. L. Chapel was starting for 
Arizona, we showed him illustrations of Hyla wrightorum. On July 6, he 
wrote from Winslow, Ariz., somewhat as follows: "At present I am at Los 
Burros CCC Camp near McNary. The last few days we have had considerable 



TREE FROGS: HYLIDAE 365 

rain. Toads are quite numerous about camp. Today we were working at Los 
Burros, which is about six miles from Winslow. In a swampy place, I heard 
quite a chorus. I found endless numbers of green frogs. No other species could 
I find. I am sending these, eight of them. . . ." On arrival, they proved to be 
four males and four females of Hyla wrightomm. 

July 9, 1942. Put up at Lake-O- Woods Lodge, Ariz. Went down to swampy 
area below the swimming pool dam. Presently in water 2-4 or 6 inches deep 
and among grass found tadpoles of Hyla wrightomm. The tails are quite 
spotted with blackish specks and in general they are rather attractive tadpoles; 
some with two legs, one or two about to have forelegs break through. When 
did they breed? 

July n, Lake-O- Woods Lodge. Today Anna and I went along a drainage 
ditch west of the lower pond and down to a swamplike area below a swim- 
ming pool. Took a lot more tadpoles ot Hyla wrightorum. 

Authorities 9 corner: 

R. Kellogg, Gen., 1932, p. 168. E. H. Taylor, Kans., 1938, p. 439. 

F. W. King, Ariz., 1932, p. 99. W. L Chapel, Ariz., 1939, pp. 225-227. 



FAMILY LEPTODACTYLIDAE 

Genus ELEUTHERODACTYLUS Dumfril and Bibron 

Map 25 

Mexican Cliff Frog, Robber Frog, Barking Frog 

Eleutherodactylus aitgitsti (Duges). Plate LXXVII. 

Range: Guanajuato. Probably not in the United States. In 1933 we gave its 
range as Jalisco to southern Arizona on the basis of the following quotation: 

"A frog of the genus Eleutherodactylus has been received from Mr. Sam 
Davidson, of Fort Worth, Texas. This specimen collected by Mr. Davidson, 
October i, 1927, in Madcra Canyon, Santa Rita Mountains, Arizona. Miss 
Doris Cochran, of the United States National Museum, to whom this speci- 
men was sent for examination and comparison, very kindly allowed Dr. Rem- 
ington Kellogg to examine it, and it was referred by him, on account of the 
peculiar dorsal spotting, to Eleutherodactylus augusti, a Mexican species liv- 
ing in the state of Guanajuato, Mexico, rather than to Eleutherodactylus la- 
trans, known only from the state of Texas" (J. R. Slevin, Ariz., 1931, p. 140). 

In 1942 Mr. Slevin showed us this specimen and we concluded that super- 
ficially it appeared to be E. latrans. After we saw Mulaik's material of young 
with light crossbands from Kerrville, Tex., we more than ever held this view. 
Koster's (Carlsbad, N. Mex.) and our (Boerne, Tex.) material also show the 
light transverse band. (Plate LXXX.) 

Habitat: (See "Authorities' corner.'*) 

Size: Adults, 2%~3 inches. 64.5-75.0 mm. (Kellogg). 

General appearance: "Mocquard concluded that E. augusti was identical 
with Cope's E. latrans from central Texas. Direct comparison of Mexican 
specimens with the cotypes of E. latrans does not entirely confirm this assump- 
tion. Though there are no constant structural features that will distinguish 
specimens from these two areas, it was observed that in E. latrans the fourth 
toe is relatively longer, the color pattern consists of fairly closely aggregated 
large black blotches, the sides and hinder half of the abdomen are faintly areo- 
late, and the skin on the upper parts of old adults is stiff, coarse, and areolate. 
These two forms are unquestionably rather closely related. An immature in- 
dividual from Jalisco and an adult individual (with a body length of 75 mm.) 
collected by Ruthling, which unfortunately is without any definite locality, 
were used in these direct comparisons. The skin on the upper parts of the im- 
mature individual from Jalisco is much more tubercular and warty than that 



kOBBER FROGS: LEPTODACTYL1DAE 367 

on the Texas specimens. Juvenile characters, such as vestigial postcephalic in- 
tratympanic dermal fold and vomcrine teeth in minute clusters, are not un- 
usual, but the presence of an abdominal disk seems rather remarkable for so 
young an individual" (R. Kellogg, Gen., 1932, p. 101). 

"Hylodes Augusti, A. Duges, notes manuscript. Aspect heavy. The head is 
so large that the body is very short; the eyes large and prominent. The tym- 
panum is conspicuously smaller than the eye. The tongue is a little longer 
than broad with a slight notch in the rear. Vomerinc teeth in 2 oblique groups 
in the rear of internal nares. The body is finely roughened on back, smooth on 
lower surface. Coloration: The upper parts are a very clear yellow, inferior 
parts white; the lower side of the throat is marbled with pale brown, the sides 
and the arms are bordered with brown. The top of head and shoulders are 
brown black, lined with pale yellow. One sees across the back a clearly defined 
band spotted with brown. This hylodes is considered very rare. M. Duges dis- 
covered it at Guanajuato, Mexico . . ." (Brocchi T.,' Gen., 1879, pp. 19-24). 

Journal notes: June 23, 1934. In the afternoon after a rain, Anna and I went 
to Madera Canyon, Ariz. Here, Mrs. Dusenberry said Mr. Sam Davidson of 
Fort Worth, Tex., stayed with her most of one summer. A woman came from 
California Academy of Science to collect. (Miss McLellan, whom he later 
married.) They collected the frog about October i. It is the specimen now in 
Calif. Acad. Sci. 

June 26, Madera Canyon. Started for old Baldy about 7:30 A.M. Arrived at 
the lookout about 1 130. L. N. Sprung of Sonoita, Ari/ona, is the fire watch- 
man. He was here in 1925 with his cousin, Kenneth Putnam, also of Sonoita. 
Last July, just below the watch station, Mr. Sprung one afternoon espied a 
queer frog on a rock face. He showed it to two boys who bothered it to sec 
it hop. It was "more round than any other shape," and after hopping on the 
rock face, it went into a crevice. It was probably the robber frog. 

In 1942 we measured the California Academy of Science specimen from 
Arizona. The measurements (in mm.) of this female arc: length 85, head 
(angle to mouth) 22, width of head 57, snout 15, eye 14.5, intcrorbital space 
5.5, upper eyelid 8, tympanum 6, intcrtympanic space 25.5, internasal space 
7, fore limb 46, first finger 15, second finger 15, third finger 2$, fourth finger 
21, hind limb from axil 106, tibia 38.5, foot with tarsus 50, foot without tarsus 
32, first toe 9.5, second toe 15, third toe 21.5, fourth toe 32, fifth toe 21.5, hind 
limb from vent 114. Forearm much developed, brachmm diameter 6, antc- 
brachium 9; suggestion of a disk on venter; has fold of skin across head from 
tympanum and one on side of body from rear of tympanum to groin, there 
bending inward to the vent. It is E. latrans. 

Authorities 1 corner: 
P. Brocchi, Gen., 1879, p. 53. 
F. Mocquard, Gen., 1899, pp. 160-163. 
R. Kellogg, Gen., 1932, p. 101. 



368 HANDBOOK OF FROGS AND TOADS 

In 1938 Taylor (Gen., 1938, pp. 391-394) described a new form, E. cactorum, 
and contrasted it with E. latrans, E. augustt, and . laticeps. He had a key 
difference: "Tympanum % to % diameter of eye; dorsal surface smooth; 
Texas; 90 mm. . . . . latrans Cope. Tympanum scarcely more than one half 
diameter of eye; Guanajuata; 75 mm. augustt Duges." In addition he gave 
a table of measurements for the three species. 

Texan Cliff Frog, Barking Frog, Robber Frog, Rock Frog 
Elctttherodactylus latrans (Cope). Plates LXXIX, LXXX; Map 25. 

Range: Waco to Helotcs to Del Rio, Tex., into Coahuila (Schmidt and 
Smith, 1944) through New Mexico to Santa Rita Mountains, Ariz. We had 
reports of it at Devil's River, Mertzon, Hondo, and Frio, Tex. R. D. Quillin 
reported it from Segovia, Medina Canyon, Boerne, Bracken, San Marcos, 
Concan, and Leakey, Tex. 

"Eight specimens from Sacaton, five miles south of Cuatro Cunegas. All 
were caught in mouse traps. Mr. Marsh states that search for specimens by 
day and night failed to discover them. They apparently got into his traps just 
before dawn" (K. Schmidt and D. W. Owens, Tex., 1944, p. 100). 

W. Chapel tells us (1945) that he caught a good-sized frog with a promi- 
nent ventral disk which he thought must be E. latrans. Alas, he lost it. He 
found it in Parker Canyon, northeast of Roosevelt Reservoir in central Ari- 
zona. It is worthy of note that there are limestone outcrops in that region. 

Habitat: Limestone ledges of the cliffs that front the Edwards Plateau. 
They have also been reported in caves, under stones and other cover, in rock 
walls of canyons, rock masses in mountains, or rocky hillsides. 

Charles T. Vorhies wrote us of his experience in Madcra Canyon, Arizona, 
"I am still sure of one thing and that is that I have never heard such notes 
from any other species of frog or toad. However, the specimen I captured was 
not on a cliff but merely on a steep rocky hillside and was sitting under a j ul- 
ting rock." 

Size: Adults, i%-3 % inches; 48-90 mm. 

General appearance: This is a large, short-bodied, squat frog with extremely 
powerful forearms and long outer fingers. Its fingers and toes have expanded 
tips and prominent tubercles on the under surfaces. The loreal region is 
oblique. The skin is smooth. The back may be light drab to greenish. 

The young and half-grown may be greenish on the back and have a light 
fawn, unspotted, transverse band across the middle back. 

Color: Male. Helotes, Tex., Feb. 21, 1925. Background of back light drab 
shading into pale drab-gray. Light area down middle of back is pale drab- 
gray or pale smoke gray. Spots on back and upper parts of arms and legs are 
natal brown or army brown; sides and sides of belly vinaceous-buflf; legs and 
arms, hands, and toes with light vinaceous-fawn or pale vinaceous-fawn; tym- 



ROBBER FROGS: LEPTODACTYLIDAE 



369 




Plate LXXVII. Eleutherodactylus au- 
gusti. 1,3. After Brocchi (X%)- 2. After 
Brocchi (XO- 4- After Mocquard 



Plate LXXV11L Eleutherodactylus ri- 
cordii planirostris. 1-4,6. Adults (X 1 )- 5>7- 
Embryo developing within the egg (X5)* 
[No free tadpole.! 8. Larva freed from 
gelatinous envelope, 3 days older than 
fig. 7. 9. Female (Xi%)- (57~9- Paintings 
by R. F. Decker t). 



370 HANDBOOK OF FROGS AND TOADS 

panum vinaccous-slate with a ventral spot from center up of pale smoke gray 
or pale drab-gray; color on rear of thigh unicolor and is the color of back- 
ground. Middle of throat and ventral parts are pale pinkish buff. Eyelid is 
Varley's gray; iris kght yellowish olive with a bronzy cast. 

In G. W. Marnock's collection, Baylor University, no. 2022 has a prominent 
transverse band across dorsum from arm insertion to arm insertion. Back of it 
comes a broad transverse area of olive-buff outlined behind by two dark spots, 
one either side of the meson. This prominent light band has a median light 
dorsal stripe extending forward on meson to between the eyes. USNM no. 
13633 (G. W. Marnock, Helotes, Tex.), which measures 48 mm., has an indi- 
cation of this median line between the eyes, and possesses a rather spotted back. 

Half-grown frog. Cave Without a Name, Boerne, Tex., from V. L. Cock- 
rell, Aug. 14, 1942. General color of snout region, area across shoulders, and 
rear of back is deep mouse gray heavily spotted with large spots of dark 
quaker drab. The area between the eyes is vinaceous-fawn with 3 or 4 promi- 
nent black spots. Across the middle back ts a light area of fawn with no spots. 
The tympanum is natal brown. The femur has 3 bands, tibia 3, metatarsus 2, 
antebrachium i of army brown, the interspaces being avellancous. The for- 
ward half of the venter is pure white with solid pigments; the lower belly, 
underside of femur and tibia, and inner side of metatarsus have no pigment 
and the flesh showing through gives appearance of light vinaceous-lilac. In 
the eye is a thin pupil rim of mars orange, the base a deep mouse gray almost 
obscured by dots of buff-yellow. 

Structure: Male with forearm best developed, brachium little developed; 
two solar, two palmar tubercles; head wide and flat; muzzle projects beyond 
lower jaw; vomerinc teeth in two short raised patches between the inner nos- 
trils; tympanum slightly deeper than wide; tongue slightly nicked behind; 
fold of skin from eye over tympanum almost to arm insertion; central abdo- 
men surrounded by a circular fold of skin; disks transverse. 

The enlarged forearm may come with age. Strecker's no. 2022 (36 mm.) 
has the forearm not so proportionally larger than brachium as it is in 74-mm. 
and 77-mm. specimens. This same specimen has not the ventral disk. 

In some alcoholic specimens there seems to be a fold of skin across top of 
head between the tympani. The fold bows slightly forward. 

Voice: At a distance the call was certainly a bark, but as we climbed the hill 
and came near, it was more of a throaty whttrr. It is ventnloquial, and the 
location of the calling frog back under the ledge doubtless increases the vol- 
ume of sound. 

"Roy [R. D. Quillin] returned Sunday [April 23, 1933] from a trip in the 
Frio Canyon [Tex.], and the barking frogs do bark. Mr. Fisher on the ranch 
said that the frogs made so much barking four weeks ago that he was awak- 
ened nights. We hear them every evening when we sit on the front porch of 
the ranch house" (letter from Mrs. Ellen Schulz Quillm). 



372 HANDBOOK OF FROGS AND TOADS 

Mr, Marnock told Mr. Cope, "During the winter the adults are very noisy, 
the rocks resounding in the evening with their dog-like bark. The noise is 
supposed by the country people to be made by lizards, especially the Gcrrhono- 
tus inferndis which occurs in the same region" (E. D. Cope, Tex., 1878, 
p. 186). 

In 1925 this last notion was still the prevalent one in Helotes, for we were 
assured that it was the "barking lizard" that made that noise. And though we 
caught the frog that was "barking" and showed it to the doubters, we judge 
many are "of the same opinion still," for since our return we have received a 
newspaper clipping with a picture and a note about the "barking lizard." 

Feb. 20, 1925. Heard "barking" near the store at Helotes. Mr. Fuller, who 
keeps the store, said that for the last week they have been calling but very 
little and slightly before that. Tonight, we went to Psettdacris ornata fork. Air 
at camp 7:45 P.M. 71; no ornate chorus frogs calling; in fact, few meadow 
frogs calling. Went on to Helotes. At the flat crossing right near Helotes ham- 
let heard a "barking lizard" or "frog." Started north along the creek, soon dis- 
covered the frog was way up on the hill though it sounded nearer at first. 
Went across open field back of the store, across fence, and up the hill among 
cedars, rhus, agerita, prickly pear, and yucca. After we thought we were near, 
we discovered it farther on. It is not startlingly ventriloquial. We began to 
suspect it was going to be in a shrubby thicket. When we neared it, it did not 
croak 5-15 times in succession but only once or twice at a time with longer in- 
tervals between croaking. At last we thought we had found the ledge. I de- 
termined to come from the left and Anna from the right of a small clump. 
I halted suddenly. Four or five feet ahead was a nice diamondback rattler 
asleep or coiled. I was searching frogs. I went around to the other side. Then 
a long interval. Then Anna thought she heard it below. Then I went above 
where I thought it was. Finally it croaked between us. We converged only to 
hear it in another direction. At last we determined it must be under or near a 
rock I was on. Anna below tried to see it, and I from above. We were about 
to give up, when I used the flashlight on it and saw it about a foot under a 
thick, overhanging rock. Could see only its pulsating throat. Anna put flash- 
light on one side to prevent its escape and with ruler poked it out far enough 
for me to get a hold on a leg. Then the game was ours. It truly is a "robber* 
frog." 

A card from A. V. Rutherford of Garner State Park north of Uvalde, Tex., 
written March 22, 1947 states: "A man here has caught them barking high up 
on the hills under ledges of rock. He heard the first barking this year, March 

15" 

Breeding: Probably they breed during any rainy period from February to 
May. It is likely that the large eggs are laid in moist or rain-filled cracks or 
crevices or even caves in the rocky cliffs and ledges where they live. 

In Sept., 1930, we saw a large female (USNM no. 10058) which was col- 



ROBBER FROGS: LEPTODACTYLIDAE 



373 



lected by G. W. Marnock at Helotes, Tex. Dr. Kellogg had discovered that it 
was a gravid female. It measures 92 mm. in body length. The egg complement 
we roughly estimate at 50 eggs. These eggs have the largest vitelli of any frog 
eggs of the United States. Measurements of four vitelli are 6.0, 7.0, 7.0, and 7.5 
mm. (/4-Kc inch). 

It seems probable to us that development is in the egg as with E. ricordii, 
yet it is possible that this large Elentherodactylns may have a method diverse 
from that small form. We were in Texas in a very dry winter and spring, but 
if the calling frogs bred, we could not find their tadpoles in the creeks, nor 
could we find pools of water near the ledges that might harbor them. 

The likelihood of their laying their eggs in a stream is about as remote as 
it is with Camp's, Marjiock's, and Ricord s frogs. (See P. strecfyri, "Authori- 
ties' corner," p. 280.) 

Journal notes: So few field experiences are described in the records of this 
frog that we are here quoting our notes at length: 

Feb. 20, 1925. Helotes, Tex. The instant I had it in hand it swelled up taut. 
When I reached the store I realized I was sweating; started to brush sweat out 
of my left eye, it smarted afterwards. Some cuts on hands also did. It affected 
mucus of mouth slightly. It certainly has a secretion. It can bloat out bladder- 
like more than any other form I have ever encountered, as much so or more 
than Hyla gratiosa or some Scaphioptts species. 

Feb. 2i. Heard about 4:^0 P.M., air temp. 71, one robber frog. It was in a 
crevice of limestone on Helotes Hill. Was near it, too. Just back of the first 
locality of last night is a Spanish bayonet. These frogs are in the limestone 
ridges. At 9 P.M., heard none. Air 70. Brec'/y but not humid. 

Feb. 22. In photographing male 1L latrans, it would wedge itself in the cell 
and climb right out of the jar. When 1 had it in hand, it bent the last joint of 
the two outer digits. These have expanded tips. With these bent digits, it can 
pull readily and strongly. It must use these for pulling itself along or to pre- 
vent animals from pulling it out. 

March 18. Tonight about 9 P.M., temp. 70, heard one calling on Helotes 
Hill, near where we heard one on Feb. 21. Why the long interval? 

March 20. Tonight about 9 P.M. heard one. Temp, about 65 F. 

March 21. Tonight after a small shower, didn't hear a robber frog on the 
hills, but at 9:30 P.M. our captive spoke up for the first time since he was a 
captive. Why? 

May 4. Last night back of camp we heard several calling until midnight. 
Air 64. John Barrara (Mexican ranch hand) said they called vigorously at 
5 P.M. We heard several north of Lee's, also east of Lee's. We heard none on 
Marnock's hill about 8 P.M. We rode almost to Babcock Road, but heard few 
beyond the area already mentioned. We heard one on hill north of our camp. 

.May 5. On hill to east heard robber frog a few times. On Marnock's hill, 
heard two or three, then they ceased calling. Not so vigorous tonight as last 



374 HANDBOOK OF FROGS AND TOADS 

night. John Barrara says they get down to the creek. The one on our hill we 
thought was perhaps moving to the creek. 

May 9. On half moon of rocks and hills east of us we heard many robber 
frogs, after the rain, about i P.M. Air temp, was 67?. This morning at 9, 1 
went on hill. Worked around top, saw a Holbroolya. Then came back at top 
of limestone ledge, and then back at base of limestone ledge, then down the 
canyon in the half moon. I did not see a water pocket of any permanence. 

July 4. Storekeeper at Sheffield, Tex., says there are no "barking lizards" 
here. About 85 miles northeast at Mcrtzon, they occur. Another man at store 
said they are also near Hondo. This accords well with information from man 
at Barton Springs Park. We were told of their being at Leakey, Tex., in the 
Frio River canyon. 

June 22, 1930, San Antonio, Tex. At night Roy Quillin and I went to John 
Classen's ranch to listen for barking frogs where he and Ellen Schulz Quillin 
hear them often. Apparently too late in year or else not rainy enough. 

June 23, Helotes, Tex. Stopped at Mr. Fuller's store. He informs me he 
heard many on the hill in the spring but not lately. 

Feb. 26, 1942. Went to California Academy; met J. R. Slevin. Glanced at 
Eleuthcrodactylus attgttsti. It's E. latrans. 

Aug. ;. Went south of Kerrville, ideal country for robber frogs, especially 
about a mile or less north of Raven Ranch where Stanley Mulaik got a few 
small ones. 

These young arc very interesting as the light transverse band is conspicu- 
ous, and their gray color in preservative is suggestive of green in life. 

Later in day went to Cave Without a Name, Mr. Cockrell guide. His wife 
said, "Surely there are salamanders and frogs in the cave. When they first 
opened it there were many black salamanders on the walls near the entrance, 
probably our cats have gotten most of them." When Mr. Cockrell came up, 
we went in. He said he saw one or two frogs on his last trip, that at times there 
were three or tour near the entrance. In the morning near the entrance he 
found one of them in a water pail. 

We started down and soon in a round pocket in the wall (pocket 6-7 inches 
deep) was a beautiful E. latmns headed outward. What a fine sight and how 
nicely niched for its life! Caught it. This one has a suggestion of the light band 
across the back. 

Authorities 9 corner: 
J. K. Strccker, Jr., Tex., I9o8c, p. 59. 
J. K. Streckcr, Jr., Tex., 1922, p. 15. 

"Enclosed is a photo of a frog I collected about 12 miles northwest of Carls- 
bad, N.M. I can't find any description in our library that fits it really well. For 
one thing the coloration is quite different. It comes closest to the Elcuthero- 
dactylns latran* collected by Marnock which you describe on pp. 161-163 of 



ROBBER FROGS: LEPTODACTYLIDAE 



375 




Plate LXXIX. Eleutherodactylus latrans. 
i. Male (X^)- 2. Male (X%). 3- Male 



Plate LXXX. Eleutherodactylus latrans. 
Upper, i. Young adult, from W, J. Koster, 
near Carlsbad, N.M. Center. 2,3. Young, 
from Stanley Mulaik, near Kerrville, Tex. 
r, 4. Adult from Cave, Boerne, Tex. 



376 HANDBOOK OF FROGS AND TOADS 

your Handbook. The following is a description that I made after the speci- 
men had been in formalin for two days : 

"Most of the dorsum greenish, darkest on head and neck, venter plain 
white. Upper jaw whitish with vertical green bars. A double whitish line ex- 
tends across the anterior part of the interorbital region. An irregular whitish 
line extends most of the distance between the interorbital line and the inter- 
scapular region. Body with numerous dark green spots, the more posterior 
ones faintly outlined with whitish. Band across back between arm insertions 
and forelimbs, ivory. Forelimbs with a bar on the wrist, forearm and a faint 
one on the upper arm. Hind limbs lighter green than the back, with darker 
green bars, these more or less outlined with lighter. 

"The colors were not greatly changed from the condition in life except for 
intensity. It was strikingly green and ivory or white when first captured. . . . 

"The specimen had an eventful history. It was taken from under our tent 
where it and a Cnemidophorus apparently sought shelter from a very heavy 
and violent rainstorm (3.48 inches at El Paso and the storm was still con- 
tinuing when we broke camp) which lasted all night and through the morn- 
ing. I heard short barks which I now suspect were these frogs but which I 
then thought were dogs. I brought it back alive and when I recovered I tried 
to identify the beast. I kept it in a deep pan of water to keep it from jumping 
whilst I was looking at the books and the darn thing nearly drowned. As a 
matter of fact it passed out and I gave it artificial respiration. It died during 
the night. After it was preserved, I set the jar on my work table next to the 
wall. A janitor moved the table for the first time in three years, broke the jar 
and threw everything into the basket. When I returned to the office I found 
the specimen in the basket with purple "Ditto" ink spots on it. It has since 
recovered. You see why I did not send the specimen right off" (letter of W. J. 
Koster, Albuquerque, N. Mex., Dec. 29, 1944). See condensed account by 
W. J. Koster, N. Mex., 1946, p. 173. 

"Though it's too long for a letter, I'll tell you my story of what I have for 
several years supposed to have been Eleutherodactylns. In 1918, while engaged 
in much field work on the Santa Rita Experimental Range, I had my family 
in a cabin well up on the east side of Madera (White House) Canyon, whence 
I came down to the range below each day. This was the last week in June. The* 
summer rains began in the canyon that week, with an afternoon sprinkle, 
next day an afternoon light shower, and next day with a rather heavy but brief 
late-afternoon rain. About dusk a loud, raucous calling broke out in the can- 
yon. I was quite new to this country, especially to summer conditions, and a 
suggestion that it was 'toads' found no favor with me. I was mystified. I went 
down into the canyon, only to find the sounds came from the opposite side; 
climbing up, all the noise hushed up. I seated myself about where I judged 
the sounds to have come from. After several fruitless minutes of waiting, I 
was startled by a sudden 'bark' directly behind me. Creeping toward an over- 



ROBBER FROGS: LEPTODACTYLIDAE 



377 



hanging rock, there was just enough light left for me to see a strange 'toad,' 
as I then thought it was. I captured it, but had no preservative, and I lost the 
specimen by escape or spoilage, I do not know which. Years later, when I first 
noted a report of Eleutherodactylus in Madera Canyon, I guessed I had seen 
and heard one. I still believe it was, and have hoped to run across it again. 
But I have never since been in Madera under similar circumstances, nor have 
I ever heard the same thing under other conditions when up there, 

"Last summer, having some field work out that way, I made a point of being 
up there on a late June day when both clouds and calendar indicated a pos- 
sible first shower, but no luck. Again, after the rains had begun, I explored 
pools along the creek in the same section of canyon without result. I'll get 'em 
yet. 

"The most I have seen of the Sierra Ancha has been in late fall no time 
to find these frogs. But the conditions would seem to be as good as in the Santa 
Ritas, barring its being further north. In fact, there are better-watered, more 
precipitous canyons there than in the Santa Ritas. Do you think I had Eleu- 
therodactylus* (letter of C. T. Vorhies, Jan., 1946) ? 

Ricord's Frog, Cricket Toad, Bahaman Tree Frog, Pink-snouted Frog 

Eleutherodactylus rtcordii planirostris (Cope). Plate LXXVIII; Map 25. 

Range: Bahama Islands, Cuba, and Florida, north to Duval County. In the 
USNM is a lot of two specimens (nos. 59419-20) taken at Auburndale, Fla., 
by Wm. Palmer. In recent years it has been taken as far north as Gainesville, 
Fla., by O. C. Van Hyning. Each of these records reminds one of Dr. Bar- 
bour's statement: "It has been taken in Florida and seems to be spreading 
northward" (Fla., p. 100). In 1931 Dr. H. A. Pilsbry, while collecting pul- 
monate gastropod Liguus on or about Long Pine Key, collected several 
(F. Harper, Gen., 1935). In 1940 W. G. Lynn and in 1943 Lynn and J. N. 
Dent spoke of its introduction into Jamaica, where it is now widespread. In 
1959 G. A. Skermer of Tampa, Fla., published some of his experiences with 
the frog. In 1944 C. J. Goin had specimens from Jacksonville. 

Habitat: Leaf piles in hammocks; rock piles; fern boxes; under walks, logs, 
debris, trash piles; limestone heaps; wells; irrigation banks. Dr. H. A. Pilsbry 
(1931) found them under loose bark in the hammocks. 

Size: Adults, %-iVr, inches (15-30 mm.). 

General appearance: It is small, elongate, similar to Syrrhophus marnoc^ii 
but possesses two long transverse curved series of vomerine teeth, behind the 
internal nares. These are delicate little frogs with the head as broad as any 
part of the body. The eyes are prominent, beadlike, bronzy, and black. The 
snout is prominent, truncate, extending slightly beyond the lower jaw. The 
legs are long and slender. The fingers and toes are very slender with terminal 
disks and prominent palettes at the joints of fingers and toes, giving a saw- 



378 HANDBOOK OF FROGS AND TOADS 

tooth appearance in latenil aspect. The color pattern varies, but in each the 
coral pink snout is prominent. 

Color: Adult. Top of snout, stripe from eye to hind leg insertion along dor- 
solateral region, interspaces of hind legs and forelegs, and the tubercles along 
the side and top of back are light coral red to dragon's-blood red or terra cotta. 
On top of snout is a dark area which joins with the dark area of the dorsum, 
which normally starts about on level with a line drawn from middle of one 
eye to middle of other. This dorsal color is brownish olive, snuff brown. In- 
distinct bars of the same color on femur, tibia, and forearm or these bars and 
the markings on the toes may be black. The upper jaw and also the lower 
jaw are spotted with alternating black or carob brown or warm blackish 
brown spots. A line of the same color over tympanum to arm insertion. This 
line sometimes continuing from arm insertion along side as more or less 
broken dark line. Sometimes from axilla to groin there is an irregular line or 
row of dark spots parallel with the upper line just described. Front of femur 
with two or three prominent black spots and several such spots on the front 
of femorotibial articulation. A horizontal line of same color from arm inser- 
tion halfway down the brachium. The under parts are white and heavily 
punctate with fine brownish olive dots. A mid-ventral blood vessel plainly 
shows through the skin of the belly. The alternating areas on upper and lower 
jaws are white. The tympanum may be almost white or with a wash of coral 
red. In front of, below, and behind eye, below tympanum, and along sides 
are spots of greenish yellow. Iris is principally coral red to dragon's-blood red 
with vermiculations of darker color. The iris rim is almost scarlet or scarlet- 
red. 

Young. The conspicuous thing about the young one seems to be the light 
coral red triangular snout patch ahead of the eye, the same color on fore and 
hind legs and a patch of same along either side. In most of the young there 
seems to be an appearance of triangular spot between the eyes, the base of the 
triangle extends between the eyes, and may be truncate, marginate, or concave. 
The rear portion of the triangle oftentimes and generally merges with the 
dorsal band, which may be scalloped along its sides. It is a beautiful little 
creature with its alternation of dark and light spots on upper and lower jaw 
and with its big bronzy eye. The legs appear barred. There is a slight ridge ofc 
skin down the middle back from near tip of snout to vent. The skin of 
upper surface is roughened and is areolate on belly and hind legs. 

Structure: Subgular vocal sac; tympanum % size of eye; tongue elongate 
oval, slightly nicked; heel reaches orbit or even snout; lower jaw with a me- 
dian tubercle fitting into a median notch in upper jaw; two metacarpal and 
two metatarsal tubercles. 

"Head as wide or wider than the body, longer than broad; the lateral out- 
lines curved; the end of the muzzle abruptly truncated. Ostia pharyngea oval. 



ROBBER FROGS: LEPTODACTYLIDAE 



379 



Vomerine teeth in two long curved series, which commence behind and oppo- 
site to the external border of inner nares, they are separated by a considerable 
space medially. Tongue elongate oval, slightly nicked. A subgular vocal sack. 
Tympanum half the size of the eye. Skin smooth above and below; sides ru- 
gose. Heel reaching the orbit. Digital palettes small. Two metacarpal, two 
metatarsal tubercles. Brachium longer than or equal to antebrachium. . . . 
A single specimen from Key West, Florida, is in the National Museum. Its 
proper habitat is Cuba" (E. D. Cope, Gen., 1889, p. 318). 

Voice: "Its twittering call can be heard from hammocks as well as dry pine 
land, after showers during April, May, June, July, and August" (R. F. Deck- 
ert, Fla., 1921, p. 22). "Their call, which they gave at night from the terrarium 
was a faint 'put put' " (E. R. Dunn, Fla., 1926, p. 155). "Their chirping notes 
are a common sound after dark and on cloudy days" (O. C. Van Hyning, Fla., 
1935, p. 4). "Some five years ago my attention was attracted by a trilling sound 
which seemed to come from a cluster of ferns around a palm. At first I thought 
it was some new cricket or grasshopper, but very careful search and watching 
did not solve the problem. Although I have lived in south Florida for forty 
years, I had not heard this trill before" (G. A. Skermer, Fla., 1939, p. 107). 

Breeding: This species breeds from April to August. Development goes on 
within the egg to adult form, there being no tadpole stage. The newly hatched 
young are %-% inch (9.0-11.5 mm.) in size. 

"Ricord's Frog does not go to the rain-pools in numbers, as do the other 
Salicntia. Pairing seems to take place on land during rainy weather, in dark 
places. The writer has so far failed to find specimens in copula, but on May 16 
two batches of eggs, containing a dozen each, were found in a depression 
filled with dead leaves and leaf-mold in a 'hammock' " (R. F. Deckert, Fla., 
1921, p. 23). 

"Ricordii lays 19-25 eggs in vegetable debris in woods, . . . yolk being about 
2 mm. in diameter, and the outer envelope eventually reaches 4 mm. No trace 
of external gills could be made out in either species. . . . The night of July 28 
a batch of 25 eggs was laid. I left Soledad on July 30 and returned on August 2 
and found that a second batch of 19 eggs had been laid in my absence. 

"The eggs of the 28th began to hatch August 7, making a period of eleven 
days. By August n all were out, making a period of ten days from the 2nd. 

"On August IT, 1925, 1 found 21 eggs in a fallen Bromeliad. These hatched 
six days after (August 16) and as soon as the yolk was fully absorbed and the 
adult coloration assumed they were seen to be ricordii' (E. R. Dunn, Fla., 
1926, p. 155). 

"Presuming that this frog, like others, laid its eggs in a lily pond near its 
habitat, I did not give thought to the life cycle. While looking over some or- 
chids which I had banked with sphagnum I found a cluster of what appeared 
to be frog or toad eggs. I had not seen eggs laid in this way before. In order 



380 HANDBOOK OF FROGS AND TOADS 

to see what had laid them I lifted the cluster, together with the sphagnum on 
which they were deposited, and placed them in a glass bowl and covered the 
bowl with a sheet of glass. 

"It was easy to see that the eggs were normal, for the embryo could be seen 
moving in them. In about seven days the eggs had hatched, the jelly-like cases 
lying flattened out on the sphagnum. After very close inspection of the sphag- 
num through the glass I saw a tiny frog not as large as a house fly seated on 
the moss. Its appearance was exactly that of the mature frog I had previously 
found so I knew that it was the young of that frog. I have found many nests 
since then, all made on the ground or close to it in damp shady places" (G. A. 
Skermer, Fla., 1939, pp. 107-108). 

"Mrs. A. N, Dow [of Jacksonville, Fla.] found eight 'small white eggs' in 
a flower pot in her yard and kept them until they hatched. The baby frogs 
were sent to me for identification, and proved to be ricordii. 

"The eight individuals vary in length from 4.5 to 5.5 mm. From subadults 
from Gainesville they differ only in having the tympanum slightly less dis- 
tinct, the head relatively broader, and the vomerine teeth less well developed. 
The eight specimens are now catalogued as number 22557 in the collection of 
the Carnegie Museum. 

"Mrs. Dow informs me that the locality where these frogs were found is in 
a residential section about 3 miles from the business center of the city. She 
found the eggs in a flower pot which had been left undisturbed in a flower 
bed in the yard for several months. The eggs were deposited on the surface 
of the soil and when Mrs. Dow first discovered them she placed them and the 
accompanying soil in a jar where they remained until they hatched about a 
week later. She sent them to me on August 2, 1945, when the frogs were 'three 
or four days old.' In answer to my query about adults, she stated that she had 
not noticed them about, but that a neighbor had seen frogs of this description 
behind the houses in the neighborhood" (C. J. Coin, Fla., 1944, P- *9 2 )' 

Apparent newly hatched young have been taken in the months of June, 
July, and August. The two following records are of young apparently not 
long after hatching: one specimen (USNM no. 36851) taken by Prof. C. H. 
Eigenmann on July 3, 1905, at Santiago de las Vegas, Cuba, measures 9 mm. 
and others (USNM nos. 29823, 29825, 29838) taken by Wm. Palmer, June 30,' 
1902, at Baracoa, Cuba, measure 11.5, n, and 11.5 mm., respectively. In the 
same lot were some specimens 15-20 mm. in length, and others 27-30 mm. in 
length. 

We have seen another lot (UMZ 64267) from Limestone Ridge, Soledad 
Cienfuegas, Cuba, 7, 7.5, 9, n, and 19 mm. in length. The 7 and 7.5 specimens 
must be near hatching size. 

Journal notes: (We have been shown the actual boards, rocks, and vegetal 
cover under which they lived and the actual flower pots in which their eggs 
have been taken, yet we have very few journal notes worthy of mention.) 



ROBBER FROGS: LEPTODACTYLIDAE 381 

Authorities 9 corner: "In the pine country [in Dade Co., Fla.] the writer 
has found it under heaps of limestone. No matter how dry the surrounding 
land may be, in the center of these rock-heaps there is always quite some 
moisture, and all kinds of creatures find hiding places there. Many specimens 
of Ricord's Frog were secured in rock pits under quite small stones. They are 
difficult to capture as they leap with lightning rapidity the instant they are 
uncovered" (R. F. Deckert, Fla., 1921, pp. 22-23). 

"E. ricordu is frequently found [in Cuba] along with cuneatus, never, how- 
ever, jumping into the water. Il is also found at considerable distances from 
the water. 

On July 25, 1924, 1 caught seven from a colony of about a dozen, which was 
under two large stones and the fallen leaves of a royal palm, and was some 
fifty yards from a stream. I kept five of these alive in a jar with some vegetable 
debris" (E. R. Dunn, Fla., 1926, p. 155). 

"Eleiitherodactvlns ncordn is a Cuban species which, it has been assumed 
(Lynn, 1940), has been introduced accidentally into Jamaica at a relatively 
recent date. It was first recorded from Jamaica by Lynn (1957) who took it at 
Montego Bay. Later, E. A. Chapm found the species at Hope Gardens in 
Kingston and it was supposed (Lynn, 11)40) that this indicated two separate 
points of introduction. The present collections however, show that the species 
is wide-spread m the island and it may be that it has been in Jamaica for a long 
time. We took specimens at Sandy Bay, Hanover, at Highgate, St. Mary; near 
Port Antonio, Portland at Chapelton, Clarendon and Mr. L. V. Burns has 
recently taken the species at Hector's River on the border between Portland 
and St. Thomas" (W. G. Lynn and J. N. Dent, Gen., 194$, p. 259). 

"Since Eleutherodactylits ricordu was first reported in southern Florida by 
Cope (Bull. U.S. Nat. Mus., (i), [875: 31) it has steadily extended its range 
northward. Carr (Univ. Fla. Biol. Sci. Ser., Ill, (i), 1940: 63) gives its range 
as continuous in south Florida northward to the region of Melbourne on the 
east coast and Clearwater on the west coast with a separate population at 
Gainesville. 1 do not know how rapidly the southern Florida population is 
spreading northward, but the Gainesville population is certainly expanding 
steadily and the species is now quite common several miles from the city in 
localities where it was not known to occur a few years ago" (C. J. Coin, Fla., 
1944, p. 192). 

Shreve's revision (Fla., 1945) suggests E. planirortris for this introduced 
frog. 

"I therefore restrict the name ricordii to the form from the highlands of 
Oriente, Cuba, which has a spotted dorsum. The color is: Above, ground 
color whitish or brownish white, heavily spotted with dark brown, the spots 
rounded and frequently coalescing, those on limbs in general smaller. In some, 
the spots coalesce to such an extent as to give the impression of marbling. . . . 

"For the form widespread in Cuba and the Bahamas hitherto called ricordii, 



382 HANDBOOK OF FROGS AND TOADS 

the name Eleutherodactylus planirostris (Cope) is available. Its type locality 
is New Providence Island, Bahamas. It is planirostris therefore, that is now 
present in Florida as an introduction. On account of the nature of the differ- 
ences existing between ricordii and planirostris (as used here), it seems better 
to use the two names in subspecific relationship, although in the material at 
hand there is no real evidence of intergradation or hybridization. This may 
be shown to occur eventually as there is some evidence of hybridization be- 
tween planirostris and the species casparit, of the Trinidad Mts. in Cuba, 
where their ranges meet, a seemingly parallel situation" (B. Shreve, Fla., 1945, 
p. 117), 

Coleman J. Coin (Fla., 1947, PP- T ~^) has just issued his excellent compre- 
hensive monograph, "Studies on the Life History of Eleutherodactylus ri- 
cordii planirostris (Cope) in Florida." It emphasizes particularly habits, 
breeding, development, color pattern, and distribution. 



Genus LEPTODACTYLUS Fitzinger 
Map 25 

White-lipped Frog, White-jawed Frog, White-jawed Robber Frog 

Leptodactyhts lubialis (Cope). Plate LXXXII; Map 25. 

Range: Mexico to extreme southern Texas in Starr and Hidalgo counties. 
First found in United States by Dr. E. H. Taylor and J. S. Wright in 1931 
(Taylor, Tex., 1932). In 1933 anc * 1934 it was held to be Leptodactylus albila- 
bris (Gunther) (Plate LXXXI) and much was made of its occurrence in 
Puerto Rico and on the mainland. Dr. Noble in 1918 pointed out three char- 
acters of difference between the mainland and insular forms but frankly ad- 
mitted that no criterion is constant. K. P. Schmidt in 1928 called the main- 
land and insular forms very close but was not able to state that they are iden- 
tical. 

Habitat: Moist meadows; irrigated fields; drains; gutters in towns; beneath 
stones and logs. Near streams; irrigation ditches; roadside ditches; in bur- 
rows in sandbanks and fields. 

"The specimen was taken late at night hopping about on the table-like top 
of one of the row of hills just north of the Los Olmos bridge, near Rio Grande 
City, Tex. The earth was dry, and no rain had fallen for a period of forty-eight 
hours. The species probably shelters itself in the crevices of the rocky ledge 
which surrounds the top of the hill" (E. H. Taylor, Tex., 1932, p. 244). 

Size: Adults, i%-2 inches (35-49 mm.). 

General appearance: This is a small dark-colored frog with a flattened 
pointed head, a low narrow lateral fold and a cream-colored labial line. 
"Among the collections made in Texas during the summer of 1931 by J. S. 
Wright and myself is a small leptodactilid toad, which offers something of a 
problepi in the way of identification. Through the kindness of Mrs. Helen T. 
Gaige, Museum of Zoology, Ann Arbor, the specimen was examined and dis- 
sected, and found to belong to the genus Leptodactyhts. She also forwarded 
for comparison specimens of the species L. albilabris Gunther from Vera 
Cruz, and St. Croix Island, which seem to approach this form in many char- 
acters. 

"From these two forms, which differ considerably from each other, I note 
that the Texas specimen differs in the absence of a distinct dorsolateral fold, 
the very much larger choanae (nearly twice the area of those in the compared 



384 HANDBOOK OF FROGS AND TOADS 

specimens), the presence of larger triangular bony prominences, on which the 
vomerine teeth occur, and their separation by a much greater distance (more 
than a half of their transverse length) ; the very narrow lateral fold is not con- 
tinuous with the small supratympanic fold. There is a small forward exten- 
sion of the snout to form a 'nose.' The dorsal coloration in life is ash gray to 
clay without black spots, while the compared forms are dark blackish brown 
with well-defined large black spots. 

"From the specimen from St. Croix (the type locality of Leptodactylus albi- 
labris Gunther) I note that the tips of the digits are distinctly less widened, 
in fact not at all dilated; that the head is longer, more pointed; the eye rela- 
tively smaller; and that the ventral brown spotting is lacking. 

"From the Vera Cruz specimens it differs in having a flatter, more pointed 
head; a different coloration. It also differs in the points mentioned above. It 
agrees with both forms in the presence of a well-defined cream-colored labial 
line, and the presence of a discoidal disk formed by a skin fold on the abdo- 
men. 

"Cope has described a species, L. labialis, which has small choanae, and a 
paucity of dermal folds, and L. gracilis with a projecting muzzle and having 
dermal folds. 

"Brocchi has described L. fragtlis with the vomerine teeth in arched series, 
with the tympanum nearly twice as large as the eye. 

"Remington Kellogg, who is reviewing the Amphibians of Mexico, pro- 
poses to place labialis Cope, and fragtlis Brocchi all under the synonymy of 
L. albilabris Gunther. 

"So for the present it seems that this Texas specimen must rest as Lepto- 
dactylus albtlabris Gunther until a sufficient series of specimens arc available 
to determine whether the separable characters, apparent in the specimen, are 
typical or due to individual variation" (E. H. Taylor, Tex., 1932, pp. 243-244). 

Color: "Above rather lead gray with a few irregularly scattered, slightly 
dark, grayish blotches; a dark vitta from snout through nostril to eye, con- 
tinued behind to the tympanum; another irregular black line begins above 
tympanum and follows the supratympanic fold; below this a narrow, well- 
defined cream line, bordered below on lip by an irregular, less intense, dark 
line continued to angle of jaw; a few black spots on posterior part of lower 
jaw; arm with a few small dark spots, and a few also in axillary region; an- 
terior side of femur with a dark horizontal line and small spots; upper and 
posterior surface of limb spotted; an indistinct cream horizontal line borders 
the granular edge of the under part of the femur, below which are a few small 
dark spots; under surface of chin through abdomen and limbs uniformly 
cream; a dark broad line from heel across foot. The measurements are, head 
and body, 25 mm.; foreleg, 13 mm.; hindleg, 40 mm." (E. H. Taylor, Tex., 
1932, p. 245). 

Structure: Cope's description of this form follows: "Vomerine teeth in 



ROBBER FROGS: LEPTODACTYL1DAE 



385 




Plate LXXXl. Leptodactylus dbilabris. 
i. After Brocchi (X%)- * After Boulenger 
(X%). 3,4- After Schmidt (X%)- 



Plate LXXXII. Leptoaaiiyim w- 
(X%)- Adults, from Stanley Mulaik, Ed- 
inburg, Tex. 



3 86 HANDBOOK OF FROGS AND TOADS 

transverse series behind the posterior border of the internal nares; toes with- 
out dermal border; no abdominal discoidal fold; posterior limbs short; end of 
metatarsus just reaching muzzle; muzzle short, not projecting; teeth behind 
choanae; one dermal fold on each side; skin rough; below white. . . . This 
small species belongs to that division of the genus, in which the toes do not 
possess dermal margin, and there is no discoidal fold of the abdominal integu- 
ment. Among these it is distinguished by the shortness of series of vomerine 
teeth and the paucity of dermal plicae. The muzzle is acuminate and rather 
narrow, but not projecting as in C. gracihs; the canthus is not distinct. The 
tongue is oval and a little notched behind; the choanae are small. The diame- 
ter of the tympanic disc is one half that of the orbit. The heel only reaches 
the orbit. The toes are not very long; there are two small tarsal tubercles and 
a narrow tarsal fold. . . . Color chocolate brown, the limbs darker cross- 
barred. A brilliant white band extends from the anterior part of the upper lip, 
and describing a curve upwards, bounds the orbit below and descends to the 
canthus oris, from which point it continues in a straight line to the humerus, 
and ceases. Inferior surfaces pure white. Length of head and body, .020; of 
head, .007; of hind limbs, 0.28; of hind foot, .013" (E. D. Cope, Gen., 1877, 
p. 90). 

Taylor states: "The snout oval, the outline broken by a slight pointed pro- 
jection of the 'nose'; the head measures, length 11.5 mm. to angle of the jaw; 
the width at same point is 9.7 mm.; canthus rostralis rounded, the ridge con- 
tinued downward in front ot eye; between this ridge and the nostril the loreal 
region is distinctly concave; eye moderate; length of orbit 3.7 mm., the dis- 
tance from the nostril 2.8 mm. and its distance from the tip of the snout 4.2 
mm., tympanum very distinct, rounded, 1.7 mm. long by 1.5 mm. high, sepa- 
rated from eye by half its greatest diameter; a slight supratympanic fold from 
eye, involving upper edge of the tympanum but bending down some distance 
behind the posterior edge of tympanum, labial fold distinct posteriorly, be- 
comes very indistinct anteriorly and terminates below nostril, in lateral pro- 
file the snout extends beyond the mouth ; the angle of mouth reaches directly 
below tympanum; the distance between orbits 2.4 mm.; of upper eyelid width 
1.8 mm. 

"Skin on head smooth, showing microscopic corrugation; the dorsolateral 
fold wanting, represented by a few low rounded or somewhat elongate pus- 
tules; a sharply defined, very narrow lateral fold extends from above arm to 
groin; above this a few elongate or rounded pustules low and indistinct; below 
this fold the skin on chin, breast, abdomen and limbs smooth except for a 
strongly granular distinctly limited area on ventral femoral region; two prom- 
inent granules below anus; a transverse fold across the breast between inser- 
tion of arms more or less connected with a lateral abdominal fold, which 
curves across the posterior abdomen, forming a disk; a small diagonal fold 
running back from edge of disk to pubic region; limbs not especially long, 



ROBBER FROGS: LEPTODACTYLIDAE 387 

the tibia 12.5 mm. is distinctly longer than the femur 10.1 mm.; from heel to 
tip of longest toe 19 mm.; an elongate sharply defined inner metatarsal tuber- 
cle, and a smaller rounded outer; subarticular tubercles strongly defined save 
under the last distal joint; heel tubercular with a distinct fold; rows of small 
tubercles below the metatarsals; tips of digits not dilated, the terminal pha- 
lanx (bone) pointed, somewhat clawlike; no webs present or only vestiges; 
two large palmar tubercles; an elongate inner at base of thumb, and a larger 
rounded basal tubercle with an anterior extension; first row of subarticular 
tubercles strongly defined; the first finger longer than second by the length 
of the last phalanx. . . . The measurements are, head and body, 25 mm.; fore- 
leg, 13 mm., hindleg, 40 mm. 

"The openings of the choanae are well back from tip of snout 2.5 mm. 
their entire outline plainly visible; vomerine teeth on two triangular raised 
patches, separated by slightly more than half their greatest width; they are 
between and almost wholly behind choanae and border the raised eye sockets 1 ' 
(E. H. Taylor, Tex., 1932, pp. 244-245). 

Voice: "The call of the male was repeated at about one second intervals 
at the height of rapidity, and resembled the plunk-plunk of a drop of water 
falling from a cave roof into a quiet pool below" (S. Mulaik, Tex., 1937, p. 73). 

Breeding: "The finding of tadpoles, transforming young and a frothy egg 
mass on June 8, 1935, immediately following heavy rains ten miles northwest 
of Edinburg, Texas, closed some gaps in the knowledge of the life history of 
this amphibian. The egg mass containing 86 yellow eggs was found in a 
rounded excavation about 4 cm. in diameter and 3 cm. deep at the base of a 
grass hummock a foot from the water edge and several inches higher than the 
water level. 

"These eggs, around which no hyaline sheath could be discerned, measured 
1.5 mm. in diameter. Within 40 hours after the eggs were laid, the young 
hatched. And were practically invisible except for two dark eye spots and the 
yellow yolk sack. Several hours later light brown pigmentation appeared. 
The emerging tadpoles measured (in mm.) 6.6 in total length, 2.3 in body 
length and 1.5 in width. Their external gills remained approximately 20 hours 
after emergence. They were apparently two branches arising near the same 
spot; the anterior branch was the larger, measuring about i mm. in length. 

"The newly hatched young rested much on their sides or even on their 
backs on the bottom of the aquaria and every few minutes indulged in vio- 
lent squirmings which brought them toward the water surface. Twenty-four 
hours after hatching, the tadpoles measured 8.1 mm. in total length, and more 
pigmentation was noted. On the second day the young were observed to feed 
apparently on algae and diatoms, and their measurements were as follows 
(in mm.) : total length 9.3; body length 3.3; body width 2. 

"On the third day measurements of u mm. in total length were recorded 
and in general the growth appeared rapid. The intestinal coil was now clearly 



3 88 HANDBOOK OF FROGS AND TOADS 

visible through the transparent abdominal wall. The animal food provided 
was not accepted before the eighth day at which time the tadpoles began to 
feed upon termites offered. Later on, as they developed, the tadpoles readily 
fed upon other insects and crushed snails. 

"On July 9 the front legs of one of the tadpoles emerged. The total length 
of this specimen was 32 mm. or 6 to 8 mm. less than the same measurement 
for field specimens taken at this stage. The tail decreased rapidly from this 
date and was gone by July n, making the tadpole stage about thirty days. 
These transforming young averaged 16.1 mm. from snout to vent; rear leg 
25.1 mm.; hind foot 12.6 mm.; rear leg adflcxcd brings heel to rear edge of 
eye; heel to tip of first toe 6.6 mm.; snout to angle of mouth 4.8 mm.; tym- 
panum 2 1 /2 times into eye. 

"Since numerous transforming young were also found on June 7, and since 
the first previous heavy rain for many months was about May 2, the tadpole 
period for the field specimens was judged to be about 30 to 55 days, about the 
same as for the aquarium raised specimens. These young were hopping about 
in the grass at the edge of the roadside ditch and were difficult to catch because 
of their agility. Though adults seemed quite common, they were more difficult 
to take. Males when calling from their cavities beneath hummocks of grass, 
left upon discovery and quickly sought shelter elsewhere. It is believed that 
the males constructed these cavities to be used by the females for the deposi- 
tion of the eggs" (S. Mulaik, Tex., 1937, pp. 72-73). 

Journal notes: May i, 1934, Edmburg, Tex. Mulaik showed me two frogs 
which are queer. These he said he found northwest of Edinburg. They were 
under trash and mud. By digging under it they found the frog. In the grass 
near by they found a female. They sounded like sea lions. 

May }. Trip with Stanley and Dorothv Mulaik northwest of Edinburg, 
Tex. We stopped beside the road near a ditch or pit where there were many 
grass clumps. Here earlier, when there was some water in the pit, they had 
heard the "sea lion" call of I^eptodactylns. In one clump they had found the 
male. Today we dug vigorously but it was too dry and no frogs appeared. 

Authorities 9 corner: 
K. P. Schmidt, Gen., 1928, pp. 38-39. 
S. Mulaik, Tex., 1937, pp. 72-73. 



Genus SYRRHOPHUS Cope 



Map 25 

Camp's Frog 

Syrrhophus cum pi Stejncgcr. Plate LXXX1II; Map 25. 

Range: Brownsville, Tex. Lower Rio (irande Valley. 

Habitat: In moist earth under hoard pile, brick pile, stones, or similar 
shelter. 

Size: Adults, r 's-i inch (15.0-25.5 mm.). 

The largest we have seen is a ripe female 25.5 mm. A collection of i; adults 
taken April 28, 1925, in Mr. Camp's board-pile station range from 15-24 mm., 
the average 19 mm., the mode 19 mm. A series of 9 taken by Mr. ('amp 
(USNM nos. 52^72-80) range from i<V-2$ mm. Another series of ID specimens 
taken by Mr. Camp (MCZ nos. 10277-86) range from 15-25 mm. 

TABLE OF MKASURKMKNTS (MM.) 



No. 

St\ 

Length 

1 lead to tympanum 

Head to angle of mouth 

Width of head 

Snout 

I-.yc 

Intcrorhital space 

Upper eyelid 

Tympanum 

Intertympanic space 

Internasal space 

Forchmb 

ist finger 

2nd finger 

$rd finger 

4th finger 

Hind limb 



Syrrhophus 


Syirhophas 




cam ft 


mainoit(ti 


USNM 


C.U. 


C.U. 


C.U. 


C.U. 


52291 


1277 


H7<> 


1271 


1382 






Ripe 9 


<J 




2O.O 


22.5 


25.5 


22.5 


26.0 


8.0 


8.0 


gjo 


<>-5 


10.5 


7-S 


6.5 


7.0 


8.0 


8.0 


7-5 


7- 1 ? 


9-5 


8.0 


9.0 


4.0 


$-5 


4.0 


4-5 


5.0 


4.0 


4.0 


4.0 


4-5 


3-5 


3.0 


? 


i-5 


3-5 


I* 


2.0 


2.O 


2.0 


2-5 


2-5 


2.5 


2-5 


2-5 


2.0 


2.0 


7.0 


6.5 


7-5 


7-5 


7-5 


2.0 


2-5 


2-5 


2-5 


3.0 


13.0 


'?-5 


'3-5 


>5-5 


17.0 


M 


3.0 


3.0 


3.0 


*-5 


2-5 


3-5 


3-5 


3-S 


4.0 


4.0 


5.0 


5-5 


6.0 


6.0 


2-5 


4-5 


4-5 


4-5 


5.0 


27-5 


31.0 


31.0 


31.0 


34-o 



390 HANDBOOK OF FROGS AND TOADS 

Sytrhophus Syt rhophus 

campi matnoclyi 

USNM C.U. C.U. C.U. C.U. 

Tibu TO.O 10.0 11.5 12.0 12.5 

Foot with tarsus 14.0 13.0 16.0 15.0 14.5 

Foot without tarsus 9.5 9.0 10.0 8.5 7.5 

rst toe 3.0 2.5 3.0 2.5 2.5 

2nd toe 4.5 4.0 4.5 4.0 4.0 

?rd toe 6 o 6.0 7.5 6.0 7.0 

4th toe 9.5 9.0 10.0 8.5 7.5 

5th toe 5.5 5.5 6.0 5.0 6.0 * 

* I hnd foot in rather poor sh.ipc. 

General appearance: This foim is very similar to Syrrhophtis marnoclyi, 
,uxl belongs to a Mexican genus. It is a gravish olive frog, with scattered dark 
spots on the back, with a dark band from the nostril through the eye, and 
with dark crossbars on the legs. The skin is finely granular. The nose is 
pointed. 

Color: Hrownsville, lex., April 2S, 1^25. Upper parts grayish olive; pale 
smoke gray from eye to eye; hair brown or bcn/o brown, fuscous, or black 
vitta in tront of eye meeting fellow on snout. This line goes through the eye, 
over the tympanum, above the shoulder insertion onto body, or may be 
broken at tympanum's lear edge and continue backward as broken spots. 
These spots may go almost to giom or stop in middle of side. Sometimes on 
kick where a lateral fold normally would be is a broken longitudinal black 
line. On the back aie scattered luscotis black or hair brown spots. Femur with 
one or two obscure spots on front. In rear, unitotm light grayish olive; tibia 
with two or three rathei prominent crossbars, interspace vctiver green. On 
sides of body aie elliptic flecks of white or pallid brownish drab, which con- 
tinue to below e\e or tarthci tot ward; one or t\\o of these spots are just belou 
and behind tympanum and sometimes some on side of face are light yellow- 
gieen. Another specimen has, starting from near vent, a row of black spots 
or one line on side ot back hallway to shoulder. Hind legs burred and spotted 
on lemur and tibia \\ith black and Isabella color, which become white or pal- 
lid brownish drab spots on foot and toes. Under parts of breast back of arm 
insertion and torward part of bell\ arc solid pale yellow-green. Blood vessel 
shows thiotigh its veij middle. Lower belly and rest of under parts are pale 
piuphsh vinaceous to light purplish vinaceous. Forearm is spotted with white 
spots, brachiurn with prominent light greenish yellow and dark spots or the 
light green-\ellow may become, in some, daik olive-buff or deep olive-buff. 
A female has fe\\ if am dark spots on the back. Ins is light greenish yellow 
above and deep chrome, with one vermiculated line of brownish through it; 
in the rear it is solid black below and in front it is black with light greenish 
yellow specks; pupil elliptical. 



ROBBER FROGS: LEPTODACTYLIDAE 



39* 




Plate LXXXIII. Synhophus campi 
(X'%). 1-8- Adults - 



Plate LXXXIV. Syrrtiopnus marnw\u 
(Xi). 1-6- Adults. 



392 HANDBOOK OF FROGS AND TOADS 

Structure: Small, delicate, flat in body; a blood vessel visible down the 
middle of the belly; fingeis and toes long and slender with prominent tuber- 
cles, which viewed from the side appear saw-toothed; forearms well devel- 
oped; tips of fingers expanded and truncate; toes less so; at least one light- 
colored tubercle just back of angle of mouth, and near the lower rear margin 
of the tympanum; often a collection of tubercles in this region or an oblique 
row to the arm insertion. Wrist extends to tip of muzzle. In alcohol, the sides 
and upper parts of hind and fore limbs seem to be covered with little round 
white spots amongst the punctae. 

Voice: This frog gives a brief cricketlike chirp, but with a whistle. It is not 
a continued call, but often consists of only one or two notes which can be heard 
a few yards away. 

April 28, 1925. Early in the morning went to the board pile station. Heard 
one chirp over in the west shed, one or two more chirping in a crack in the 
brick wall near the ground. Heard several in the board pile. 

June 17, 1950, Brownsville, Tex. They are a dooryard, front-porch, or garden 
friend, cheerful as a cricket. The note may be one ttcl{ or two or three ticl(s, 
these well measured. Or the two mav be rapid and close together. Sometimes 
after these comes a cricketlike call. 

Breeding: This species breeds from April to May. The eggs arc few, 6-12, 
and very large, the egg yolks about Vis inch (3.0-3.5 mm.). Larval develop- 
ment is probably within the egg. The smallest frogs we have seen in collections 
range %-%o inch (5.0-8.5 mm.). 

April 28, 1925, at Brownsville, Tex. We secured a good series of these little 
frogs. One female contained very large eggs. 

May 5, Helotes, Tex. This evening about 5:30 P.M. I looked at the Browns- 
ville Syrrhophus cumpi to compare with our lone 5. mtirnocfyi. Lo and behold, 
a male was holding a female in inguinal amplexation. The fists met. One fin- 
ger reached across the thigh and the other across the lower belly. The male 
is more spotted; the female almost without spots. 

May 6. Tonight we had company. When I showed them the S. campi speci- 
mens, there were three in one embrace, one above another. Each of the upper 
two had an inguinal embrace. 

A ripe female taken April 28, 1925, has six or seven very large ovarian eggs 
well advanced. These egg yolks range from 5.0-3.5 mm. in diameter, mosfof 
them about 3 mm. 

The very few large yolked eggs and the very small frogs seem to confirm 
the opinion of Drs. Stcjneger and Barbour and Messrs. R. D. Camp and A. C. 
Weed that the larval development is carried through in the egg. 

A lot of 10 small frogs sent to Dr. Stejnegcr by Mr. Camp measured 6.5, 7.5, 
7.5, 9, 9,5, 9.5, TO, 10, 11.5 mm. Others equally small were seen in Mr. Camp's 
collection. The American Museum has 3 specimens (7.5-9.0 mm.). The Mu- 
seum of Comparative Zoology has 18 small specimens: one 5 mm., one 6 mm., 



ROBBER FROGS: LEPTODACTYLIDAE 



393 



six 6.5 mm., seven 7 mm., one 7.5 mm., one 8 mm., and one 8.5 mm. (5-8.5 mm.) . 

Journal notes: April 27, 1925. Mr. Camp of Brownsville, Tex., took me to 
the type locality. It was in a small back yard of a private residence in the center 
of the city. In the rear ot the house was a laundry, the floor of which was close 
to the ground. Against the rear of the house were two small piles of brick, 
each of which was no more than 2-3 feet across and i%-2 ft. high. The ground 
was moderately moist there. Mr. Camp said, "Move these bricks and you will 
find them." He warned me to grab quickly but I was not quick enough for 
the first one. I had turned over two or three small piles of bricks and become 
quite discouraged when I espied another under the shade of a little fig tree. 
Presently I caught my first one. The associates of these creatures are sow bugs, 
ants, snails, and spiders. Something jumped in a hole; all I got was the im- 
pression of the jump. In another case there was a jump under a board which 
was under the brick pile. Saw 12 or 15, but all I captured were four adults 
17 mm., 20 mm., 24 mm., 24 mm. and a half-grown one 12 mm. in length. 
Only a keen collector would have discovered this new species. 

April 28. It certainly seems as if they come out to the edge of the board pile 
at dusk or in the evening. Under one square board i ft. or i% ft. found four 
at the edge of the pile. Do they feed at night ? Someone said they did not 
want to lose them all because they fed when mosquitoes do and fed on mos- 
quitoes. 

April 29. Went to the board pile at 9:30 A.M. mighty hot. Once or twice 
while we were moving the pile, we heard the chirp of this species. Beneath this 
pile is an old cistern, now partially filled with bricks and dirt. It is very moist in 
this cistern, but no water. We saw two or three frogs in amongst the bricks. 
We moved about two-thuds of the pile, secured eight or nine frogs. We saw 
no signs of eggs. One we caught rather harshly gave up a lot of cockroach 
eggs. These he wouldn't need to feed on at night. We didn't find any of the 
very little ones, cither in the board pile or the yard. These frogs were very 
quick, leaping the instant they are exposed. They arc very flat and, like the 
robber frog, have the forearm well developed. When under board or cover, 
they often crawl along. 

June 16, 1930. Walked around Brownsville about 11-12 P.M. Was sure the 
cheerful chirp I heard in many yards was Syrrhophns campi. 

June 17. About 5 P.M. went over to the board pile station. The board pile 
which Camp, Weed, and we have successively moved and put back again 
is gone, but Mr. O. P. Hacker who owns it (corner Washington and T5th 
Street, Brownsville, Tex.) said there are frogs in the same old place. Mr. A. 
Holm tells me this area was General Zachary Taylor's old stables. Went into 
a shed near the old place. Began turning over debris, sticks, wet papers, iron 
pieces, and clothes. Under these the little frogs were revealed. The first two 
I lost. Some appeared quite "yellowish green" or old gold with spots. They 
can crawl rapidly. Most of the specimens were quite dark. In various places 



394 HANDBOOK OF FROGS AND TOADS 

while at the task I heard their pleasant notes. They are quick to find a crack 
in the wall or other cover. Where the old board pile was, now bricks are in a 
pile. These I didn't move but in the pile were frogs chirping. Took several 
small frogs. One very tiny. 

June 19. Mr. Rebb's palm grove. Arrived at 7. Palm grove flooded. In palm 
grove are plenty of S. campi near its unflooded edge. Others sound as if in 
deeply flooded parts. Are they temporarily on boles of trees, on large palm fans, 
or in a litter of palm leaves at the surface ? Surely this species is not scarce. It 
is common. 

May 2, 1934, Harlingen, Tex. Mr. and Mrs. L. Irby Davis say they have 
plenty of small frogs under their plant boxes. They are S. campi. Saw several 
of them at the edge of their rear lawn. 

Authorities 9 corner: 
L. Stejneger, Tex., 1915, p. 132. 

Gaige's Frog 

Syrrhophits gaigcae. Map 25. 

Range: "Type from the Basin, Chisos Mountains, lirewster County, Texas 
No. 27361 Field Museum of Natural History. Collected July 24, 1937, by 
Walter L. Necker. . . . 

"The sixteen paratypes agree well with the additional specimens which were 
taken in the Basin and one from Pulham Canyon by the junior author. Ten 
additional specimens from Juniper Canyon were collected by Mrs. Helen T. 
Gaige . . ." (Schmidt and Smith, Tex., 1944, P- 80). 

Habitat: After our examination of the Gaige material of the University of 
Michigan some 15 years ago we wrote: "The Gaiges | Dr. F. K. and Mrs. Helen 
Thompson] became very familiar with this species while they were collecting 
in the Chisos Mountains, Texas. They came to know its nocturnal habits and 
its notes and collected a good series of adults." How appropriate that in 1944 
the authors Schmidt and Smith "named | this frog | for Mrs. Helen T. Gaigc, 
in order to associate her name with the hcrpetological exploration of the Chisos 
Mountains, in which she had a pioneer part." 

Size: Adults 21-28 mm. Mrs. Gaige's ten, collected 15 years earlier, we made 
21-31 mm., i.e., slightly lower than our and Schmidt and Smith's range for 
S. niarnoctyi. 

General appearance: "Closely allied to Syrrhopus marnocfyi, from which it 
is distinguished by smaller size, wider head, and vermiculate instead of spotted 
dorsal pattern. . . . These range in length from 21 to 28 mm., compared to 25 
to 35 mm. in marnoclyi; the ratio of width of head to body length is 0.40 in 
seventeen gaigcae and 0.36 in twenty marnocfyt. The paratypes agree with the 
type in vermiculation and diflfusencss of the dorsal pattern" (K. P. Schmidt 
and T. F. Smith, Tex., 1944, p. 80). 



ROBBER FROGS: LEPTODACTYLIDAE 



395 



Color: "Pale brown above, with dark brown reticulation; under surfaces 
uniform very pale yellow; limb with broad dark crossbars" (same). 

Structure: "Syrrhopus gaigeae sp. nov. . . . Description of type. Habitus 
of Syrrhopus imtrnoclyt, head wider than body, its width equal to the distance 
from snout to posterior border of tympanum; snout long, the nostril close to 
the tip; canthus rounded; length of eye much less than its distance from the 
nostril, about equal to the diameter of the tympanum, which is very distinct; 
heels overlapping; heel reaching midway between eye and tip of snout; disks 
of two outer fingers much wider than those of inner fingers; disks of toes 
small; first finger about equal to second; metacarpal, metatarsal, and subdigital 
tubercles distinct; skin of back slightly tuberculated; a rounded gland above 
the insertion of the arm; no glandular fold over tympanum" (same) . 

Journal notes: We have no actual specimens. Once in the Davis Mountains 
we were sure we saw a Syrrhophus disappear under a huge stone and in the 
same way we were unsuccessful in Devil's River. Is there a wide gap of 250 
miles in Syrrhophus range or a wide gap in our knowledge? Again, Edwards 
Plateau to Trans-Pecos needs attention. 

Authorities' corner: In the account of this species we have used Schmidt and 
Smith's account almost exclusively for there is little else of record. They write 
(p. 80): 

"The new form is admittedly distinguished from marnocfyt by only trivial 
characters; it is believed, however, that the differences noted characterize a 
well isolated population, separated by some 250 miles from the range of 
mar noc kit, and that these differences indicate that a distinct form is in the 
process of evolution." 

About 1930 we measured two of the University of Michigan specimens, each 
28 mm. in length, and compared them with two S. marnocl{n of 28 mm. For 
the 25 measurements we then made, we found the Tran-Pecos specimens fell 
within the range of the 25 S, marnocfyi measurements. 

We have measured mature S. marnoclyt from 22.55 mm. to 33 mm. and two 
28 mm. S. gaigeae. We find our measurements are recorded thus: Width of 
head in body length 2.66-2.8 in the two S. gaigeae specimens within the whole 
range of 2.66-3.29 in our S. marnocfyt specimens. Or put as the dcscribcr docs 
(ratio of width of head to body length) it would be 0.357-0.375 for S. gaigeae 
and 0.30-0.375 for S. marnoctyi. The actual measurements for the width of 
head in the two S. gaigeae arc 10.5 mm. and ro mm. 

Marnock's Frog 

Syrrhophus marnoc\ii Cope. Plate LXXXIV; Map 25. 

Range: Helotes near San Antonio, San Marcos, and Austin, to San Angclo, 
Tex., and Sonora. 
Habitat: Cracks, crevices, caves in limestone ledges of the hills and ravines. 



396 HANDBOOK OF FROGS AND TOADS 

We made our captures under large flat stones near limestone ledges, one above 
the wall of a ravine. 

Size: Adults, 7 <6-i% inches (22.0-39.5 mm.). 

Our two specimens were 22.5, 26 mm. (See S. camfi for detailed measure- 
ments.) 

The American Museum has one (no. 22664) collected by Byron C. Marshall 
at Ezell's Cave, San Marcos, Tex., May 12, 1928, which is 24 mm. 

The Museum of Comparative Zoology has two specimens (nos. 18527, 
185273) from Hclotes, Tex. They measure 22 and 25 mm. 

The Baylor University Museum has G. W. Marnock's collection, some 
adults of which vary from 23-55 mm., and others taken by W. J. Williams in 
1927 and 1928. One of these is 31 mm. 

General appearance: This small, flattened frog has a long flat head, with a 
broad space between the eyes, which are large and prominent, indicating noc- 
turnal activity. In general color, the back is greenish, spotted with brown, the 
under parts light-brownish vinaceous, the legs crossbarred. The nostril is far 
in front of the eyes, which are as far apart as the breadth of the hump. The 
skin is smooth. There aie one or two white or deep colonial bufl tubercles be- 
low and behind the tympanum. 

Color: Helotcs, Tex., May 5, 1925. General color greenish or olive lake; top 
of head from forward part of upper eyelid mignonette green or grape green or 
light yellowish olive, the upper eyelid rainctte green and unspotted. In fact 
whole top of head from upper eyelid forward is unspotted and sharply marked 
oil by the vertical purplish lorcal area. The dorsum from upper eyelids back- 
ward becomes ecru-olive and the upper parts of forehmbs arc the same or 
lighter, a deep colonial bufT. On the upper surface of hind limbs this color 
changes to olivc-ocher, yellow-ocher, old gold, or yellower than any of these 
three. Spots on back, black, sorghum brown, or vinaceous-brown. The tympa- 
num and across shoulder insertion is light purplish vmaccous. The groin is 
purplish vinaccous. Underside of hind legs, forelegs, and across breast arc light 
brownish vinaccous. Throat pale grayish vmaceous; sides light brownish 
vinaccous or pale grayish vinaceous; back of eye, some of tympanum, canthus 
extending around snout, labial rims, sorghum brown; chin specked with 
sorghum brown. On sides of body are some white or pallid brownish drab or 
pale russtan blue spots; one or two white or deep colonial buff or chamois 
tubercles below and behind the tympanum. Belly, pale yellow-green or pale 
fluorite green; a purplish vinaccous blood vessel down middle of belly divides 
the color into two parts. Iris dull green-yellow above and below with sorghum 
brown vermiculatums or spots on it, or upper part \\ith one wavy longitudinal 
line and then small spots; front and rear of pupil dark vinaccous-brown, thus 
from snout to rear of eye one continuous vitta. 

In some alcoholic specimens, the upper parts from head to vent are very 
finely spotted; and in some of the smaller specimens, the following transverse 



ROBBER FROGS: LEPTODACTYLIDAE 397 

bars on the hind legs may be indicated, two on the tarsus, two on the tibia, one 
just below the knee, and two on the femur. On the tibia and femur, one of the 
two bars may be light-centered. 

Structure: Small flattened body; eyes prominent, widely separated by broad 
flat space; fingers and toes long and slender with prominent tubercles, which if 
viewed from the side give the digits a prominent saw-toothed appearance; tips 
of fingers are expanded and truncate, toes less so. Wrist of forehmb extends 
beyond extremity of muzzle. Foot 10-11 mm. Tarsus 6.5 mm. No vomcrine 
teeth. 

In several we have seen white tubercles on right and left sides of lower belly, 
also on rear of femur. Arc they natural or parasites? 

Voice: This is a cricket! ike chirp of one or two notes of an instant's duration, 
possibly followed by a trill of two or three notes. 

May 6, 1925, Helotes, Tex. While I was describing this specimen it chirped 
on three different occasions, and reminded me of the chirping S. cam pi of a 
week or more ago at Brownsville, Texas. 

We searched with flashlights more than once along the muddy bank of 
Helotes Creek at the flat crossing near Marnock's home for the source of a 
similar chirp. We thought it might belong to a frog, but search as we would, 
we could not find the musician. It was back a little from the edge of the water, 
where the mud was soft, punched by cows, and more or less covered with 
grassy tussocks. Similarly we searched in Brownsville on Camp Brown 
grounds for a chirp on the side of a grassy ditch. We had no success. 

May 10, Helotes, Tc\. On our hill, we went walking at 6 P.NI. Nighthawks 
booming; a few robber frogs calling. Below limestone ledge heard a crickctlikc 
call frequently followed by a trill of two-three notes. A froglikc note, Acru- 
like only shorter. Is it Marnock's, in .imong rocks and oaks? 

Breeding: Development is probably within the egg and during April and 
May. 

A few of the larger specimens in collections taken in May look to be ripe, 
and almost all the specimens captured have been secured in that month. We 
suspect that the month of May is an important part of the breeding period. 
Eggs not described. 

It seems likely that these frogs hatch fully developed from the egg, just as 
Dr. Stejnegcr and Dr. Thomas Barbour suggest likely for S. cumpi. 

There is probably no free-swimming tadpole and no transformation in that 
sense. There is no record of a specimen of the small si/,c (5 or 6 mm.) of 
S. campt. The smallest we have seen are two taken by W. ]. Williams. They 
are: Baylor University Museum no. 2185 taken at San Marcos, March }, 1927, 
and another, no. 3853, taken at San Marcos, June 1928. Each is 17 mm. in length 
of body. 

Journal notes: May 5, 1925, Helotes, Tex. In Helotes Creek Canyon above 
camp . . . where a side ravine comes in, we were searching for a poinsctt's 



398 HANDBOOK OF FROGS AND TOADS 

lizard we had seen. . . . About 3-4 feet farther up the side of the canyon was 
a large, flat, loose stone on a stony slope. I lifted it and a small frog hopped 
out. I got the impression of a greenish frog. I dropped the stone quickly to see 
where the leaping frog went but lost it. Then I lifted the stone again, and at 
first my eye did not espy the second frog. Had it been a Camp's frog it would 
have leaped. Presently my eye saw a Syrrhophus. It is Marnock's frog. It is 
bigger than Camp's, but much like it. 

Our second frog was caught May 10 on Marnock's hill, back of the store at 
4 P.M. In ihe morning it rained hard until i P.M.; then we went to Marnock's 
hill. East of the store one-half mile, found on top of the hill a pebbly place 
with a few flat stones. A banded gecko and Marnock's frog were under a stone. 
It was moist, black dirt beneath the stone, and it was in a more or less open 
place with dwarf oaks about. 

June 23, 1930, San Antonio. I left at 8 o'clock for Helotes with Gable and his 
son, Hugh. They parked their car at a crossing above G. W. Marnock's house. 
They started up a horseshoe hill I never was on in 1917 or 1925. 1 suspect this 
was Marnock's special hill for collecting. His house is in the opening of the 
horseshoe. After we reached the top, we went down toward a glen where there 
had been seepage earlier. In this place they had found two Marnock's frogs, 
which they gave to Mr. Parks to send to General Biological Supply. One of 
these very frogs we received this spring. Their habitat was much the same 
as that of those found by us. 

Authorities 9 corner: 
W. J. Williams, Tex., 1927, p. 7. 
}. C. Marr, Gen., 1944, p. 480. 



FAMILY RANIDAE 

Genus RAN A Linne 
Maps 26-36 

Texas Gopher Frog, Southern Crayfish Frog, Southern Gopher Frog 

Rana areolata areolata Baird and Girard. Plate LXXXV; Map 26, 

Range: Matagorda Co., Tex., north to McCurtain Co., Oklahoma, and La- 
fayette Co., Ark.; probably also in extreme northwestern Louisiana (L. Stej- 
neger and T. Barbour, Check List, 1943, p. 53). 

Habitat: "I saw but one specimen of this frog and collected that on June 
1 8 at the Machine Gun Range, twelve miles out from Camp | Houston, Tex. |. 
It was a half-grown frog and was hiding Under a log at the edge of a pool" 
(P. H. Pope, Tex., 1919, pp. 97-98). 

Size: Medium, length 2 1 /2-?H inches (62-90 mm.). 

General appearance: This is a stocky, medium to large, rather rough- 
skinned, brownish frog with round dark spots on back and sides, with reticula- 
tions of dotted light lines and with prominent dorsolateral folds. 

"Rana areolata. . . . Appears to be closely allied to/?, halecina \pipiens\ and 
utricularia, but the head is larger and the spots of the back are smaller and more 
numerous. Male with an external vocal vesicle on each side behind the angle 
of the mouth . . ." (G. A. Boulenger, Gen., 1882, p. 41). 

According to Cope (Gen., 1889, p. 409), R. v. areolata has "length of head 
to posterior tympana three times in total; tympanic disk round; dorsal spots 
well separated; nostril equidistant between end of muzzle and eye," whereas 
R,. v. circulosa has "length of head one third of total; tympanic disk variable, 
dorsal spots so large as to leave only circles of the light ground-color; nostril 
nearer eye than end of muzzle in young." 

Coin and Netting (Fla., 1940, p. 146) give these contrasting characters. 
"Head U-shaped in outline when viewed from above; dorsum often smooth, 
or nearly so; tibia length less than 40 mm. in adults; post-tympanic fold poorly 
developed; dorso-lateral folds narrow or only slightly raised, or both. . . . 
Rana areolata areolata Baird and Girard. Head orbiculate in outline when 
viewed above, dorsum rugose, tibia length more than 40 mm. in adults; post- 
tymphanic fold well developed; dorsolateral folds prominent. . . . Rana 
areolata circulosa Rice and Davis." 



400 HANDBOOK OF FROGS AND TOADS 

Bragg (Okla., 19423, p. 18) of Oklahoma, which state has both areolata and 
circulosa, characterizes them as follows: "Dorsal surface smooth; dorsolateral 
folds narrow and usually not well developed; legs fairly short (tibiae 40 mm. 
or less). Distribution, southeastern Oklahoma. Rana areolata areolata, South- 
ern Crayfish Frog. Dorsal surface often rugose; dorsolateral folds well de- 
veloped; leg rather long (tibiae more than 40 mm.). Distribution northeastern 
Oklahoma. Rana areolata circulosa Rice and Davis (Northern Crayfish 
Frog)." 

Color: Houston, Tex., from R. H. Vines through the kindness of A. C. 
Chandler, March 5, 1948. Three males. The background of the backs ranges 
from chalcedony yellow to olive-yellow. In the darkest one there are more 
speckles on the background, which is deep olive-buff. In two, the heads are 
much darker than the backs, and the larger spots are inconspicuous there. In 
one, the head is like the back, the spots clearly showing. The large collapsed 
vocal sacs are courge green to light-hellebore green. When one frog blew up 
his pouch to a diameter of % inch, the pattern of spots showed in a dull 
and dusky manner. The concealed surfaces of hind legs and the groin are 
suffused with citron yellow to strontian yellow. 

In the smallest and lightest frog, the background of back and head is dark 
olive buff except the snout, which is darker. On the back between dorsolateral 
folds are three irregular rows of spots of buffy olive, outlined by primrose 
yellow. On the side are three more irregular rows, but these spots are not com- 
pletely encircled with light and hence are not so conspicuous. They are light 
brownish olive on a background of vinaceous-buff to wood brown, which is 
marked with conspicuous round spots of ivory yellow. The center of the 
tympanum is marguerite yellow, ivory yellow, or white. The bars on the legs 
are very prominent, 3 on tibia and 2 on thigh with i at the knee, and are col- 
ored and outlined like the spots of the back. On the tibia each interspace be- 
tween bars bears a thin line of light brownish olive. On tarsus and outer edge 
of foot are 2 bars. The exposed surface of the femur is very spotted but the 
spots are not outlined and their color is olive-yellow. This spotted pattern is 
revealed on the venter and on the rear margin of the femur. The ventral sur- 
face of the femur is light vinaceous-fawn. The forehmbs are not conspicuously 
barred, being only irregularly spotted. The under surface is white. The edge 
of upper jaw from eye backward is mottled, and the rim of lower jaw bears 
few dark spots. There is a spot on each eyelid and one on the interorbital space. 
The eye is large and bulging, the black iris bearing fine dots of zinc orange 
with the warm buff pupil rim notched below. 

In the largest frog, the light spots of the sides are often centered with pin 
points of black and are raised warts on mounds. Similar wartlike spots cross 
the rear of the sacral region. The black pin points are so conspicuous on the 
back that this is a very speckled frog. Between the coastal folds are 5-7 low 
irregular broken rows of longitudinal folds. 



FROGS: RAN1DAE 401 

Structure: Length of head contained in total length three times. Length 
of leg to heel equals length to eye or nostril. Tibia longer than femur. 

"Rana areolata, B. & G. Head very large, sub-elliptical; snout prominent, 
nostrils situated halfway between its tips and the anterior rim of the eyes, 
which are proportionally large. The tympanum is spherical, and of medium 
size; its central portion is yellowish-white, whilst its periphery is black. The 
body is rather short and stout; the limbs well developed; the fingers and toes 
very long without being slender. The ground color of the body and head is 
yellowish-green, marked with dark brown. Besides there arc from thirty to 
fifty brown areolae; margined with a yellowish line. The upper part of the 
limbs is of the same color as the body, but instead of areolae, transverse bands 
of brown are seen on the hind ones. The lower part of the head and body is 
yellowish, with small dusky spots along the margin of the lower jaw, and 
under the neck. A specimen three inches and a half long was found at India- 
nola [Texas], and a very small one on the Rio San Pedro of the Gila" (S. F. 
Baird and C. Girard, Gen., 1852, p. 173). This last frog might well be another 
form, or did R. areolata occur in Arizona? (Note suggestion on p. 516.) 

"Skin rough, with elongated warts on back and sides. Lateral folds con- 
spicuous. Tibia ridged lengthwise. No glandular fold along the jaw. A distinct 
fold over the ear from eye to shoulder. Under and posterior surface of femur 
granulated. Head large; unusually thick through. Muzzle long; space between 
eyes greater than width of eyelid; nostrils nearer to the end of the muzzle 
than to the eye; eyes large; ear half to two-thirds size of eye. Foot with a tarsal 
fold; webs short (three joints of fourth toe free); inner sole tubercle small, 
no outer tubercle, tubercles under toe-joints prominent" (M. G. Dickerson, 
Gen., 1906, pp. 192-193). 

Breeding: February to June. 

"As regards an easy way of capturing Texas gopher frogs, there is no such 
thing as an easy way in my limited experience with the species. In fact, I cap- 
tured so few of them and those under conditions of unusual expenditure of 
energy that even at this date I can remember each individual capture. I have 
verified my memory by looking up my notes. 

"The first is a male captured under circumstances which suggested the mat- 
ing urge was upon him. The second was observed while feeding on insects 
under a street light on Main Street, just outside Hermann Park. It was cap- 
tured after a merry chase that left me rather muddy. I was returning to the 
Institute when Dr. and Mrs, Chandler stopped their automobile by the curb 
for a little conversation. The remaining frogs were all captured on February 
ii, 1930, after a careful stalk of a breeding pool where the frogs were singing. 
In this case the stalk was performed so slowly that more than an hour was cov- 
ered in approaching the pool where three clasping pairs were taken. The fe- 
males extruded their eggs that night. Incidentally these frogs secreted a foamy, 
sour-smelling material that made my hands itch. 



4 02 HANDBOOK OF FROGS AND TOADS 

"I think the only means of capturing these frogs with any assurance of suc- 
cess would be to stalk the animals in the breeding pools. I made many unsuc- 
cessful attempts to capture these frogs before I approached sufficiently slowly 
and carefully to get them under the flash light. On land these frogs are very 
active. My experience suggesting they are even more difficult to capture on 
land than the southern leopard frog, but in the water they are very slow to 
move and can be seized easily. However, they are very shy and if one ap- 
proaches the breeding pool at a normal pace, these frogs are nowhere to be 
seen when one arrives. 

"The pool where I caught the mating pairs was located northwest of the 
chemistry building, but all that portion of the campus is now in cultivation 
and I dare say that the pool has been destroyed; the frogs dispersed to other 
areas. Breeding pools are easily located by the mating song, which is produced 
after each heavy rain in late January and February. The mating song is easily 
learned, because so far as I could tell it is exactly like the mating song of the 
most common toad of Houston, namely, Bufo valhceps. However, the toad 
sings in May and June, while the gopher frog sings in the winter" (letter from 
P. D. Harwood to A. C. Chandler, June 12, 1946). 

Journal notes: Northeast of Edinburg, Tex., Aug. 3, 1942. Ten or fifteen 
miles north of Raymonclville a dead Rana pipiens or areolata in the road. 
Mesquite on either side. A many-spotted frog we could see, and little else. 
Another frog so crushed I could not tell it. 

Authorities* corner: 
M. C. Dickerson, Gen., 1906, p. 193. 
C. E. Burt and M. D. Burt, Tex., 19293, p. 6. 
A. N. Bragg and C. C. Smith, Okla., 1943, p. 107. 

Northern Gopher Frog, Gopher Frog, Crayfish Frog, Hoosier Frog, 
Crawfish Frog, Ring Frog 

Rana areolata ctrculosa Davis. Plate LXXXVI ; Map 26. 

Range: "From Rogers and Tulsa counties, Oklahoma, north through east- 
ern Kansas, eastward across central Missouri and Illinois to Bcnton and Mon- 
roe counties, Indiana (possibly to Greene county, Ohio), and southward in the 
Mississippi Valley through western Kentucky and Tennessee to Pontotoc 4 
County, Mississippi" (L. Stejacger and T. Barbour, Check List, 1943, P- 54)- 

Habitat: Old burrows, commonly crayfish burrows in the vicinity of ponds. 

In 1910, "Professor La Rue found the frogs in the mammal burrows along 
the shores of the ponds, as well as in crayfish holes, but it is probable that they 
were only temporarily occupying the former during the spawning season for 
we were unable to discover any mammal burrows, either in the vicinity of 
ponds or elsewhere inhabited by frogs" (C. Thompson, 111., 1915, p. 6.) "The 
old burrows occupied by crayfish were entirely without chimneys, and were 



FROGS: RANIDAE 



43 




Plate LXXXV. Rana areolata areolata Plate LXXXVl. Kana areowiu t t ^. 

tyu\ Ma i es 1,4. Females (X ] /6)- 2. Female (X%)- 3- 

VA Male 



4 o 4 



HANDBOOK OF FROGS AND TOADS 



approximately round at the entrance, which had a diameter of about three 
inches. The entrance was more or less overhung with grass, and at one side 
was a small bare space about six inches in diameter. . . . The burrows oc- 
cupied by frogs differed but slightly from those just described" (same, p. 3). 

"The habitat is, in general, low meadowland, which is sufficiently moist to 
harbor crayfishes. The burrows of the latter are necessary, apparently, for the 
well-being of this frog, which lives in them during the day. . . . Holes with 
funnels, of course, can be inhabited by nothing but crayfish; but frequently or 




Map 26 

usually they have nothing but the smooth platform in front always in this 
case they are inhabited by frogs. These platforms serve to prevent grass from 
growing at the entrance and provide mounds upon which the frogs sit at night 
or early in the morning to cjtch stray insects" (H. M. Smith, Kans., 1934, pp. 
480-481). 

"During the postbrcedmg season ctrcttlosa resorts to crawfish burrows and, 
therefore, is not found in streams or ponds. Dr. Wilfred Crabb and Mr. George 
Wiseman collected four specimens on May 6, 1942 at Stockport, Van Buren 
County. The following habitat description is quoted from their field notes: 
'The frogs were found sitting near the openings of holes into which they re- 
treated when disturbed. They were extricated by fish hooks attached to the 
ends of slender sticks. The holes were apparently vacated crayfish burrows. 



FROGS: RANIDAE 



405 



One was three feet deep with 18 inches of water in the bottom, another was 28 
inches with 14 inches of water. Two frogs were usually present in the same 
burrow although the large one [adult male; snout to vent length 67 mm.] was 
alone. There were about 12 holes within a radius of 50 feet and all contained 
frogs. They were located on a short-grazed bluegrass pasture with a slope of 
7 per cent. The day was cloudy and cool (about 40?.) with a fresh west wind.' 
Another specimen taken in a similar situation the same day in an adjoining 
section was an adult female 101 mm. in length" (R. M. Bailey, la., 1943, p. 350). 

Size: Adults, 2^-4 % inches. Males, 63-104 mm. Females, 75-11? mm. 
Among Percy Viosca, Jr.'s material from Ottawa, Kansas, from Gloyd, were 
males 105, 108, m mm. and one 92 mm., having thumb and sacs well de- 
veloped. At Monroe, La., P. Viosca, Jr., and H. B. Chase had caught a female 
of 87 mm. (maturity not certain) and one young 46.5 mm. 

General appearance: This is a large brownish frog with round dark brown 
spots on back and sides. These spots may vary in size, as large or small. They 
are in three or four rows between and sometimes extending onto the dorso- 
lateral folds and several irregular rows or groups on the sides. The spots are 
surrounded with grayish white borders, which on the lower sides become the 
background color between the spots. The skin of the back is roughened with 
tubercles. There are one or two long glandular folds on the tibia. There is con- 
siderable greenish yellow in the groin and on the concealed portions of legs 
and feet. There is a prominent dark bar on the brachmm, a broad, fleshy, 
spotted band along the jaw, and a light center to the tympanum. The light 
color on the arms and jaw is grayish. There are several smaller, hglu-rimmccl, 
dark spots on eyelids, between the eyes and on top of the snout. The legs aic 
prominently barred with brown and cream or gray. In this group, the frogs 
have a few dark spots just forward of the arm insertion. When cold and wet, 
the frogs were very dark. The vocal sacs of the males are large and conspicuous 
when collapsed, and bluish gray-green in color. When expanded they form 
large balls on either side above the arms. (When plowed out in early spring 
they are so dark as to be almost blackish.) 

Color: Calhoun, 111., from CI. A. Nicholas, Feb. 27, iy$o. Mule. Down the 
back between dorsolatcral folds arc three rows of spots. Back of the hump 
mainly two rows. Spots mummy brown, raw umber, brownish olive, or clove 
brown. These are encircled by thin lines of ivory yellow or cartridge buff or 
marguerite yellow. This color merges into interstices between spots. These 
interstices are covered with small dots of the four browns given above for big 
spots, producing in effect a light grayish olive or grayish olive. This color on 
top of forearm becomes light mineral gray. It is grayish olive on hind legs. 
The femur, tibia, and tarsus have mummy brown or one of the other three 
browns in their conspicuous crossbars. The interspaces between bars are like 
interstices of back, but with small partial crossbars of the darker color. On 
sides are at least two rows of large spots; mummy brown bar on forearm in- 



406 HANDBOOK OF FROGS AND TOADS 

sertion. Underside of forefeet and hind feet pale vinaceous-drab or light gray- 
ish vinaceous. Under parts, throat, breast, belly, and lower sides pale cinnamon- 
pink or cartridge bufl (before molting it was colonial buff or deep colonial 
bufl) . Same color on fore half of venter of hind legs and underside of fore- 
arms. Underside of femur, down middle, pale vinaceous-drab or light grayish 
vinaceous. Top of foot and tarsus yellow ocher or honey yellow. Wash of picric 
yellow on rear of arm, axil, groin, front of thigh, and rear of tibia. On rear of 
thighs and somewhat in front oi thighs the color becomes oil yellow. Edge of 
lower eyelid bluish gray-green or parula blue. Pupil horizontal, broken in 
middle below. Over pupil obscure longitudinal band of xanthine orange in 
front and rear or with considerable empire or pinard yellow. Pupil has rim of 
this yellow. Iris in general black. Vocal sacs tea green to celandine green. 

Female. Encircling light lines around large dorsal spots, general interstices, 
interstices of hind legs, and light spot in tympanum, pale olive-buff or tilleul 
buff, on sides becoming even vinaceous-buff. The dorsolateral fold is tilleul 
buff or vinaccous-buff. This interstitial color is with many fine dots of olive 
brown or deep olive. The large dorsal spots, three rows forward and two back- 
ward of the hump, are clove brown or fuscous. These spots are smaller on sides 
and become natal brown. The crossbars on hind legs and spots on forelimbs 
are like spots of dorsum. Spot across arm insertion in front and one just beyond 
insertion on arm in axil. Side of head, edge of lower jaw, and venter ahead of 
arm insertion with small olive-brown or clove brown spots, giving side of head 
a speckled appearance. Axil, rear of arm, groin, front of thigh, and rear of 
thigh greenish yellow or green-yellow. On reticulated rear of femur this be- 
comes oil yellow or courge green. Dark color is fuscous. Front of tibia and top 
of foot cream-buff. Under parts are pale pinkish buff or marguerite yellow. 
Thigh on underside has same light grayish vinaceous of male. Also same color 
on underside of foot and hand. Ins black or fuscous. Pupil nm vmaceous- 
tawny or vinaceous-fawn. Over pupil, a horizontal band or dash of these 
vinaceous colors. Same color in fine dots on lower iris and outer part of upper 
iris. 

Structure: Large; skin warty on back and sides; head shorter, mouth smaller, 
and hind limbs longer than R. capita; males having prominent, collapsed, 
pleated vocal sacs resting outside like folds of skin and continuing along the 
sides as folds past the axil, the middle of the sac being back of the tympanym; 
thumb somewhat enlarged in the male; eye conspicuous, but small in relation 
to snout; fourth toe very long; a thickened or fleshy band along the edge of the 
jaw. On the breast, the arm insertion with the pectoral girdle is conspicuously 
indicated by a triangle of much thinner skin, the base at arm's insertion, the 
point at the pectoral region; waist slightly broader, thus making whole form 
less wedge-shaped than R. capita. 

In Jordan's Manual of Vertebrates of the Northern United States . . . (2d 
ed., rev. and enl., Chicago, 187(8), page 355, we have this description: "R. circti- 



FROGS: RANIDAE 407 

losa, Rice and Davis (sp. nov.) Hoosier Frog. Head broad; body, head and 
sides with the ground color largely predominating, and with narrow rings of 
a greenish slate color, which becomes larger and more irregular posteriorly; 
hind legs black, crossed with irregular lines of yellowish slate color; fore limbs 
similarly marmorate; tympanum black with pale ring; below chiefly yellowish 
white; toes very long; size medium, L. 3/l>. Benton Co., Indiana, lately dis- 
covered by Mr. E. F. Shipman (abridged from Mr. Rice's Notes)." 

"There is a strong dorsolateral glandular ridge on each side, and between 
these there are from six to eight narrow glandular folds not so much broken 
up as in the R. a. aesopus, but readily becoming indistinct in alcohol. The dorso- 
lateral fold extends nearly to the groin. Below it the sides are crowded with 
longitudinal glandular folds, more or less broken up. . . . 

"Since the above was written I have been able, through the kindness of Pro- 
fessor Forbes, of the university at Champaign, 111., to examine the type speci- 
men of Messrs. Rice and Davis. It differs considerably from the specimens 
above described, as follows: The muzzle is not protuberant, so that the nostril 
is equidistant between the end of the muzzle and the eye, as in the subspecies 
arcolata. The tympanic disk is nearly round, and its long diameter is three- 
fourths that of the eye. This specimen has twice the bulk. In other respects 
it does not differ. A very strong glandular thickening of the skin extends from 
the eye above the tympanum, and then descends posterior to it. The eyelid also 
is thickened. 

"Two specimens (No. 13828) from Olney, III., also received since the above 
description was written, explain these discrepancies. The larger of the two 
agrees with the type in all respects, but the smaller, which about equals the type 
in dimensions, has the elongate muzzle of the small ones that I have described 
above. In both the tympana are three-fourths the orbit, and in neither is it de- 
cidedly oval" (E. D. Cope, Gen., 1889, pp. 41 $-415). 

Voice: "A loud trill, hoarser than that of the leopard frog and pitched some- 
what higher than that of Rana catesbeinna" (C. Thompson, Mich., icji5a, p. 6). 
To Mr. Ackert, the call sounds "half -strangled" as if it had its mouth half out 
of water. 

"The call, from near at hand, is of the general type of that of Rana palustris, 
but is much louder and quite distinct in timbre. It may best be described as a 
deep guttural, snoring sound, with a slight upward crescendo at the end. It 
is given at intervals of perhaps ten to fifteen seconds, though we did not time 
it" (H. P. Wright and G. S. Myers, Ind., 1927, p. 174). 

"The song of Rana arcolata was most often heard after dark although on 
one occasion several were singing and splashing in a roadside pond about an 
hour after sundown. The voice of these frogs does not have the prolonged 
resonance of that of the bullfrog, R. catesbeiana, although it is almost as deep 
and seems to have even more carrying power. The song most frequently heard 
is a low-pitched, drawn-out guttural note which may be suggested by the 



408 HANDBOOK OF FROGS AND TOADS 

syllables "wurr-r-r-up" accented on the last. It is repeated several times, either 
from the surface of the water or from the shore, at more or less regular inter- 
vals, varying in frequency. The vocal sacs of the males are lateral and rela- 
tively much larger than those of R. ptpicns. When singing they are distended 
until they resemble miniature balloons, each one almost as large as the head 
itself" (H. K. Gloyd, Kan., 1928, pp. 117-118). 

"A deeply sonorous and resonant 'w-a-a-ah'; loud and of exceptional carry- 
ing quality. From mixed choruses this stands out above the others; clearly 
audible for over half a mile. The strong choruses are heard in April" (R. M. 
Bailey, la., 1944, p. 17). 

Breeding: March to mid-May. The National Museum (no. 48697) has a ripe 
female of 103 mm. taken at Montgomery, Miss., on March 9, 1911, by Mr. 
Parker. (See Gloyd in "Authorities' corner.") We ourselves have not seen the 
egg masses, but they are doubtless phnthhke like those of Rana capita. 

On April 1 1, 1926, near Bloomington, IncL, Herman P. Wright and George 
S. Myers found 17 egg masses of this species in a small pond not more than 
90 feet in diameter. 

"I have found these frogs singing in numerous choruses during the last of 
March and first of April m southeastern Kansas. Near Lawrence they were 
collected while breeding on April 27, 19.51. In 1933 mated pairs were collected 
at the same locality on April 2r. Gloyd . . . states that they were last heard 
singing in 1927 in Franklin county on April 9; the first specimens were taken 
March it. ... 

"Temporary pools by roadsides and in pastures are chosen in which to breed 
and lay eggs. Males sing at the edge of the pools or out in the water, and, al- 
though they cease singing frequently upon the approach of a light, yet they 
will remain above water until splashing about or other noises cause them to 
cluck beneath the water or to sidle back into their holes. In some pools it fre- 
quently was possible to capture them after they disappeared by passing the 
hands back and forth over the mud and grass at the bottoms of pools, where 
they remain hidden until further disturbed or danger is past. A number of 
specimens placed in a tank with three or four inches of water illustrate well 
this protective instinct. Upon being startled by some sudden activity outside, 
they duck to the bottom of the water, close their eyes, push their heads down 
against the bottom of the tank, and propel themselves blindly forward by slow 
alternate or coincident strokes of the hind legs, holding the front legs against 
the body. Such actions in pools on soft earth would very quickly cover them 
from sight" (H. M. Smith, Kans., 1934, pp. 479-480). 

"Laid in large plinth-like masses about 5-6 inches in diameter in shallow 
water about stems of grass, etc. Probably about 7000 eggs are in each mass. 
Individual eggs are rather distinct. The outer membrane measures about 
4.5-5.0 mm. in diameter, the inner about 3.15, and the vitellus about 2.46-2.50. 
The vitellus is considerably larger than in pipiens, and the space between the 
inner and outer membrane is greater than in the latter species" (same, p. 479). 



FROGS: RANIDAE 409 

The transformation time is the first week of July (H. P. Wright and G. S. 
Myers, Ind., 1927, p. 174). Size, 20-30 mm. Herman P. Wright and George S. 
Myers were to have described the tadpoles. We had tadpoles of this species 
from Antioch College, Yellow Springs, but they were without eyes, etc., i.e., 
experimental specimens. A number of tadpoles secured 4 miles north of Bloom- 
ington, Ind., April 12, 1940, by H. T. Gier were loaned to us by Mittleman. 
They measured 39.5-51.0 mm., the bodies being 20-22 mm. Most of them were 
approaching transformation. Three transformed and transforming ones were 
20, 20, and 25.5 mm. 

The Museum of Comparative Zoology has two specimens (no. 1478), col- 
lected in Michigan by J. G. Shutc, 1863, which are slightly past transformation. 
They are 33 and 34.5 mm. 

"Sexual maturity is apparently attained at an age of not less than three years. 
Three early spring collections of overwintering tadpoles show no sign of ap- 
proaching metamorphosis, but since a unimodal size dispersion is evident . . . 
[a table gives overwintering tadpoles ranging from 31-63 mm.], transforma- 
tion may be assumed to occur during the ensuing summer. Three frogs taken 
by Crabb and Wiseman on May 6 were immature; they have snout to vent 
lengths of 48 ( 9 ), 49 ( 9 ), and 51 ( <J ) mm., and are judged to be two years 
old, having transformed during the preceding summer. The adult male 67 
mm. in length taken with them is presumably a year older; it is possible, 
however, that he transformed early in the summer of his first year and was 
mature at two years of age. Since this is easily the smallest adult taken it is 
believed the bulk of breeding specimens are at least four years old" (R. M. 
Bailey, la., 1943, p. 350). 

Journal notes: Sept. u, 1929. At n A.M. went with Dr. Fernandus Payne to 
the Rana areolata pond found near Bloommgton, Ind., in 1926 by H. P. Wright 
and G. Myers. It is the only pond in that vicinity in which they have found 
this frog breeding. They discovered it by hearing the frog chorus. The pond is 
Vi mile from the road. There arc cultivated fields (cornfields) on one side 
of it and meadow on another. On one side is a hedgerow of a few willows and 
oaks, and near by are a few shrubs, but in the main the pond is open. It was 
dry in the summer and the bottom mud cracked into blocks. A recent rain 
formed a small pool in the center. We could not tell where m the surrounding 
country the Rana a. circulosa would be likely to have gone. We went on to 
Olney, 111., and south to Calhoun to look at those habitats. There we looked 
up Lee Ackert, who went out as a young lad with the Thompson girls, when 
they were working on this form. He took us to their farm a mile or so east of 
Calhoun. Here in a field that contained an open pond we searched for the 
characteristic holes these frogs occupy. He said the holes were 2V&-3 inches 
across. We found nothing promising in or near the pond. The fields were so 
thickly grown up to grass and pasture weeds that we found very few holes. 
The boy dug up two or three but had no luck. He said he has plowed them 
up and cut them in plowing. Thought possibly a field not worked this year 



410 HANDBOOK OF FROGS AND TOADS 

would be better. Went to a pond a short distance north and showed us places 
there where the "girls" found them. 

Mr. Nicholas reported plowing up several in the spring. We sought him i% 
miles southeast of Calhoun. He took us down his lane where 2 or 3 weeks ago 
he saw a Ran a a. circulosa at the entrance of a 2-3-inch hole. We dug and dug; 
finally at about 4 feet we reached water level. He reached in several times and 
was nipped twice. Proved a crayfish. Did he drive out the frog? Went to house 
and field on opposite side of road. In a lower portion near woods, which he 
tells us is covered with water until late in April, we searched for likely holes. 
Found one but it was filled in. Mr. Nicholas told us that sometimes two or 
three frogs may be in one such hole with heads near the opening. Sometimes 
they appear to be in colonies when plowed up. They appear quite blackish 
when broken out and look almost the color of a catfish. They work down into 
the dirt again after being disturbed. They are large, heavy frogs, clumsy and 
slow-moving. His place is near the source of Sugar Creek. Sugar Creek Prairie, 
a known habitat, is southwest of him about 2^-3 miles toward Parkers- 
burg. 

During three warm days at the end of February, 1930, he caught and sent 
us four frogs, one male, three females, from this prairie. To secure one of these 
he told us he dug down 5 feet. 

Authorities' corner: 

G. E. Beyer, La., 1900, p. 37. J. Hurter, Mo., 1911, pp. 116-117. 

F. A. Hartman, Kans., 1907, pp. 226- H. K. Gloyd, Kans., 1928, pp. 117- 

229. 1 1 8. 

Oregon Red-legged Frog, Western Wood Frog 

Rana aurora aurora (Baird and Girarcl). Plate LXXXVII; Map 27. 

Range: British Columbia, Washington, Oregon, south along the coast of 
California to Eureka. 

Habitat: "Oregon and Washington specimens were taken among the ferns 
and dense vegetation in the forests of the coastal belt" (J. R. Slevin, B.C., 1928, 
p. 129). 

"This frog is abundant in the region of Portland throughout the year except 
in December, when all of our frogs are in hibernation. It is an inhabitant of 
the deep forests, resting under the cool sword ferns of this region during the 
day and often coming forth at night in search of food" (S. G. Jcwett, Jr., Ore., 
1936, p. 71). 

"It is usually found in the ferns and vegetation of the woods, or in nearby 
wet areas" (K. Gordon, Ore., 1939, p. 62). 

March 16, 1942, Clam Beach, Calif. In a pussy-willow pond took an 
Amt>ystomci-l\ke. mass of eggs. In a pond just back of beach dune behind a 
eucalyptus cover took several Ambystoma masses well along and one Rana 



FROGS: RANIDAE 



411 



aurora subsp. Saw two Rana aurora 
on logs and one gave a Rana catlike 
call when it leaped. They are not very 
fast but live among bad tangles of 
pussies and logs and tussocks. 

Size: Adults, r?4~3% inches. Males, 
44-63 mm. Females, 52-87 mm. Great- 
est size less than Rana aurora dray- 
tonii. Storer (Calif., 1925, p. 231) had 
males 50-58 mm. and females 57-76 
mm. in length. 

General appearance: Distinctly a 
wood frog, the mask is evident in 
many. It is medium in size, moder- 
ately stout, and smooth-skinned. The 
head is narrowly oval from above, the 
profile thin. The back is brownish or 
olive, frequently with inky spots. 
Sometimes a few of these spots have 
light centers. Frequently there is a 
dark bar across the upper arm. There 
is red on the sides of the body and on 
the concealed parts of the legs, feet, 
and underarms. The light line along 
the upper jaw ends in a fold at the 
corner of the mouth. Groin mottled. 

March 31, 1942. J. R. Slater com- 
pared three Ranas of Oregon and 
Washington. 

Mottled red, yellow, and green in 
groin; R. aurora aurora. 

Mottled somewhat in groin and 
lower sides yellowish, cream; R. cas- 
cadac. 

Not mottled in groin; groin red; 
R. prcttosa. 

April 2, Longview, Wash. Just east 
of Stella, at basaltic falls beside road, 
next the water's edge took one small 
jR. aurora aurora. At west edge of falls 
in big cavities among large rocks and 
logs took five R. aurora aurora, not a 
fast frog nor hard to catch, yet it is 





R. a. c/rayfonii 
R. cascadae 



Map 27 



4 i2 HANDBOOK OF FROGS AND TOADS 

more alert than R. pretiosa pretiosa. The groin is black marked with con- 
spicuous spots of pale greenish yellow, which area extends slightly onto the 
belly. 

"The Red-legged frog may be recognized by its slender form, smooth skin, 
dark patch behind the eye, and red colour on sides of body and under surfaces 
of legs" (G. C. Carl, B.C., 1943, p. 48). 

Color: Female. Tacoma, Wash., from Prof. James Slater, April 21, 1930. 
Underside of tibia, front half of venter to femur and along sides from groin to 
axilla solid coral red, jasper red, or light jasper red. A wash of same color on 
hind toes and membrane of web. Top of back, snout, fore limbs, hind limbs 
brownish olive, light brownish olive, Saccardo's umber to umber. Sides below 
costal fold with considerable black, area back of eye more or less black but not 
a clear vitta. Deep colonial buff or deep olive-buff or Naples yellow line from 
shoulder insertion below tympanum to below eye is broken up on side of snout 
with many black dots. Throat heavily specked with smoke gray, pale olive- 
gray, or mouse gray. Breast and belly white but with much gray (of the shades 
listed above) on it, though not so uniformly overlaid as on throat. Groin has 
black and some light dull green-yellow spots. Eye barium yellow, iris border 
above ochraceous-orange or zinc orange on lower ins rim, which is broken in 
middle. In lower iris are specks of same color as lower iris border. In upper ins 
is a little ochraceous-orange. Bars on hind legs not very distinct, black with 
many ground color specks on them. 

Male. Carbon River Valley, Rainier National Park, Wash., from J. R. Slater, 
May 16, 17, 1930. Dorsal color sulphinc yellow, citrine, dull citrine, or buffy 
citrine with few faint black specks. On brachium color becomes old gold. 
Hind legs and forelegs same color as dorsum. Stripe along upper jaw to over- 
arm insertion barium yellow to Naples yellow or cream color. Mask back of 
eye extends diagonally down to end of cream-colored upper labial stripe. Mask 
obscured by many minute oil yellow specks. Iris black with prominent lemon 
yellow or light greenish yellow bar across eye above pupil. Lower eye mainly 
black with few English red and xanthine orange spots. Upper eye with few 
specks above light greenish yellow bar. On sides more black specks. In groin 
several larger viridine green spots. Lower sides, undersides of tibia, and top of 
hind foot and femur slightly Etruscan red, old rose, Corinthian red, or deep 
vinaceous. Femur hone brown with white specks. Under parts dirty white or 
yellowish glaucous. In general venter cloudy. In the young the upper labial 
light line is particularly conspicuous. Iris golden yellow. 

Structure: Frog medium in size; skin smooth; ridge from angle of eye to 
shoulder not prominent; dorsolateral folds indistinct; webs very well de- 
veloped, greater in males than females; some males with inconspicuous groove 
across middle of thumb tubercle. 

"The general aspect of this species differs greatly from all its congeners in 
North America. The length of the body and head together is three inches and 



FROGS: RANIDAE 413 

a half, the head forming nearly one third of this length. The head itself is 
pyramidal, pointed, the nostrils situated midway between the anterior rim of 
the eye and the tip of the snout. Eyes of medium size, anterior limbs short; 
fingers rather long and slender. The body is orange red, with here and there 
black irregular patches. From Puget Sound" (S. F. Baird and C. Girard, Ore., 
1852, p. 174). 

Voice: Seldom recorded in the field. Like the wood frog it has a short breed- 
ing period so that few hear it to know it. 

Breeding: These frogs breed from February to May. Eggs are 3.04 mm., 
outer envelope is about 12 mm. The tadpoles have not been described. They 
transform at 11 /io-% inch (17-21 mm,), and the labial teeth are %. 

"We collected a large series of adult and newly transformed Rana aurora 
aurora in April 1933, ]ust north of Klamath, Del Norte County. It was raining 
and frogs were abundant on the sopping forest floor. On subsequent examina- 
tion one of the supposed red-legged frogs turned out to be a female Ascaphtis" 
(W. F. Wood, Calif., 1939, p. no). 

Wallace Wood's individuals that transformed in April interest us. We early 
recorded transformation at 17-21 mm. and observed: All this material 
(USNM no. 337) except two specimens, 26 and 44 mm., we interpret as re- 
cently transformed. The National Museum frogs were taken at the Columbia 
River on a United States exploring expedition, and Dr. Barnhart states that 
the expedition came to the Columbia River "late in April, 1841," and arrived 
two weeks later at Puget Sound. After our varied acquaintance with this form 
we conclude that these and Wood's transformecs must have come from early 
breeders. 

"Little is known of the life-history. The eggs are laid in masses during 
March and April in backwaters of lakes and streams. Tadpoles collected in a 
lake near Victoria, B.C. on May cjth, 1941, when about i inch in length, trans- 
formed in July of the same year" (G. C. Carl, B.C., 194?, p. 48). 

"Eggs: The mosaic appearing masses are made up of large eggs with a vol- 
ume of about 1.2 cc. each. The jelly is very transparent, loose, and viscid. Three 
envelopes are present in addition to the vitclhnc membrane, which is easily 
discernible in most cases. Although rather indistinct all envelopes can be seen 
without the aid of a lens when viewed in proper light. Most difficult to make 
out is the middle envelope, which is the thinnest. The fairly large vitellus is 
black at the animal pole and creamy white at the vegetal. Measurements are: 
vitellus 3.04 mm. (range 2.^1 to 3.56 mm.); vitellinc capsule 3.28 mm. (range 
2.56 to 3.87 mm.) ; inner envelope 570 mm. (range 4.0 to 6.68 mm.) ; middle 
envelope 6.80 mm. (range 6.25 to 7.93 mm.) ; outer envelope 1 1.</> mm. (range 
10.0 to 14.0 mm.). 

". . . Several differences are noticeable between the eggs of /?. a. aurora and 
its very near relation R. a. draytonii. In general, eggs of R. a. aurora are larger; 
the first thing to strike the eye, the vitellus, is perceptibly greater as is the 



414 HANDBOOK OF FROGS AND TOADS 

dimension of the outer envelope. The outer envelope is also less distinct in 
R. a. aurora. Vitelline membranes are more easily seen in R. a. draytomt. With 
R. a. draytonii the three gelatinous envelopes become progressively greater in 
thickness from the inner to the outer, in R. a. aurora the middle envelope is the 
thinnest. The entire egg mass of R. a. aurora is from two to four inches greater 
in diameter than that of R. a. draytonii. 

"R. a. aurora eggs somewhat resemble those of R. sylvattca. The looseness 
of the jelly and form of the mass are similar, but the lack of a middle envelope 
and usually smaller dimensions of individual eggs and masses readily separates 
R. sylvattca from R. a. aurora" (R. L. Livezey and A. H. Wright, Gen., 1945, 
pp. 702-703). 

Journal notes: March 27, 1942. Stopped between Hoh and Borachiel rivers, 
Olympic Peninsula, Wash., beside a wonderland meadow, with spots of snow, 
Ledum, cranberry, etc. Here in ditch saw several fresh R. a. aurora egg masses. 
Saw one composite mass of frogs' eggs getting stranded, 2 ft. by 3 ft. in 
diameter. (Must have been 15 to 30 masses here. Mating congress in one place 
as in R. sylvatica.) In each egg envelope were green algae as in salamander 
eggs- 

March 19, Oregon. From 11:00 A.M. to 12:50 stopped between Hauser and 
Lakeside. In a series of pools beside a road in a wood clearing recently cut off 
were several pussy-willow ponds. One has cattails; each has some sedge tus- 
socks. All have some log entanglements. On these logs, along the banks, and 
in shallow mounds in the pond these frogs sometimes sit, not infrequently 
two in a place, either young or females. Lots of vegetation in bottom of ponds. 
Centers sometimes 3 feet deep. No eggs. All adults caught are females. Where 
are the males? The females all ripe. Some of the young are bright red under 
legs and even on sides of pectoral region. Throats much spotted. . . . Later 
took in a ditch, i foot wide but 2 feet deep, one female at least 20 feet from 
pond, on way to pond. These frogs surely look like wood frogs, and are wood 
frogs. 

March 20. Up McKenzie R. with Ruth Hopson. . . . Caught little R. 
aurora and saw two adults. What is mass of Ambystoma-hkc eggs left high 
and dry? 

March 27. Between Nolan Cr. and Hoh River found beside road a ditch at 
least 4 feet across, containing plenty of vegetation and with sedge clumps on 
its wooded side. Beside sedge clumps are Arnbystoma-like masses. In shallow 
end of ditch Anna espied frogs' eggs, quite dark in color. The eggs are far 
apart m mass, at least % inch from each other. The fresh mass is quite bluish 
in cast, and in water 6 inches deep. One mass at least 8-10 inches across! An- 
other 6 inches across! Three good masses and a partial one; must be 
R. a. aurora. Where are the R. a. aurora males and females? Heard seven. 
Stopped beside another stream. Here found two Rana, one adult with much 
red and cloudiness and one small one, yellow below. 



FROGS: RANIDAE 



415 




Plate LXXXVII. Rana aurora aurora, i. Plate LXXXVIII. Rana aurora draytonii 

Male (X%) 2. Female (X%). 3. Female (X 1 /^). 1,2,4. Females. 3. Male. 



416 HANDBOOK OF FROGS AND TOADS 

March 30. Went to Sparaway Lake with Mr. James Slater and later to 
another more brushy lake. Here he gets R. a. aurora. Found one mass of 
JR. a. aurora eggs. The egg has two egg envelopes with a very visible vitelline 
membrane. The tadpoles have three rows of teeth on the upper labium and 
four rows on the lower. 

Authorities corner: 
H. S. Fitch, Ore., 1936, p. 640. 
I. McT. Cowan, B.C., 1939, p. 48. 
W. C. Brown and J. R. Slater, Wash., 1939, pp. 17-18. 

"Some previous collectors have placed specimens of this species [Rana cas- 
cadae\ with Rana aurora aurora. . . . Rana aurora aurora and Rana cascadae 
overlap a little in distribution in the Canadian Zone" (J. R. Slater, Wash., 
I939b,p. 149). 

California Red-legged Frog, Drayton's Frog, Bullfrog, Long-footed Frog, 

Bloody Nouns, Rocky Mountain Frog, Western Wood Frog, 

French Frog, Leconte's Frog 

Rana aurora draytonii (Baird and Girard). Plate LXXXVIII; Map 27. 

Range: Coastal California to Lower California. Few records in Sierras or 
east of them. Introduced in Nevada (Lmsdale). "Inhabits chiefly the Upper 
Sonoran life-zone, but extends into Transition and Lower Sonoran" (J. Grin- 
nell and C. L. Camp, Calif., 1917, p. 149). 

Habitat: Ponds, lakes, or marshes. Large permanent pools or water courses. 
Linsdale in Lower California (L. C., 1932, p. 353) gives several habitats. "More 
than half the specimens in the scries listed above were captured in or at the 
margins of streams. One individual was taken from a spring, one from a 
slough, one from the ground beneath willows near a creek, two in rat traps 
at the edge of a swamp, three from a marsh, and four from a lake. One of the 
frogs was captured by the use of a trout fly." 

"It is known to occur in a number of water-storage reservoirs and other 
artificial ponds, and it has been cultivated with some degree of success in ponds 
of quiet water on one or more 'frog farms' within the State. ... In general, 
the ponds and creek pools which are the haunts of this species are seldom 
frozen, and if they do freeze, the ice seldom lasts for more than part of ^ one 
day" (T. I. Storer, Calif., 1925, pp. 236-237). 

"This is the large frog which frequents the ponds and streams on both sides 
of the mountains. It is usually found where water is permanent and when 
disturbed takes refuge in the weeds at the bottom. It has been collected in 
Imperial County. It may be distinguished from the other frogs of the genus 
Rana in this territory (except the Leopard frog) by the ridges on the sides of 
the body, and from that frog by the absence of the conspicuous oval blotches 
which characterize the latter" (L. M. Klauber, Calif., i934a, p. 7). 



FROGS: RANIDAE 417 

Size: Adults, 2^-5% inches. Males, 63-100 mm. Females, 58-136 mm. 

General appearance: This is a large, stout, rough-skinned frog with a thick 
broad head. It is the largest of California's native frogs. The leg bars on tibia 
and femur may be so prominent as to appear zebrahke. The back is olive drab 
or buffy brown, spotted with light-centered spots; the groin, heavily spotted; 
the lateral fold, light pinkish cinnamon; the line on the upper jaw, cream 
buff or pale pinkish. The underside of the hind legs and the inner half of 
the tarsus and foot are pinkish to red. The young are apt to be conspicuously 
spotted. 

Color: Sentenac Canyon, San Felipe Cr., Calif., from L. M. Klauber, March 
25, 1928. Female. Upper parts Isabella color, Saccardo's olive, drab, or buffy 
brown. Lateral told light pinkish cinnamon. Line on upper jaw pale pinkish 
cinnamon or cream-buff. Brighter than in male. Around tubercles between 
lateral folds and on sides are black rings with light pinkish cinnamon in 
center on tubercle. On sides this becomes congo pink or dragon's blood red. 
Groin is dark purplish black and spotted with olive-buff or cartridge buff. 
Femur marked with dark purplish black and cartridge buff on dorsal sur- 
face. Underside of hind legs and inner half of tarsus and foot, pale pinkish 
cinnamon, congo pink to dragon's blood red. Iris orange. 

Male. Upper parts citrine-drab or yellowish olive to light grayish olive on 
forearms; more uniform on back with few black spots or specks. Groin more 
spotted than in female, more yellow present. Ventral parts more heavily 
spotted and reticulated; otherwise as in female. 

Structure: Thumb enlarged, slightly two-lobed; thumb base quite large in 
female and in some individuals would be mistaken for a male; prominent 
ridge from posterior angle of eye to shoulder; skin thick and roughened with 
many small papillae, even the eardrum may be so roughened; eardrum small; 
dorsolateral folds prominent; sacral hump prominent; back with regularly 
placed light-centered spots; male without visible vocal sac. 

"This species resembles very much ... \R. aurora\ in its external appear- 
ance. It differs, however, in having a truncated snout, the nostrils consequently 
nearer to its tip than to the eyes. The eyes themselves and tympanum are pro- 
portionally larger than in R. aurora, the limbs more developed and the tongue 
much narrower. The ground color is olivaceous green, maculated with black 
on the upper region of the body and limbs, whilst underneath the hue is uni- 
color, except sometimes under the head, breast and hind legs, where the brown 
and white mingle in circular dots. Specimens were collected at San Francisco, 
California, and on Columbia River by Mr. Drayton himself, to whom we 
take pleasure in dedicating this species" (S. F. Baird and C. Girard, Ore., 
1854, p. 174). 

Voice: "A series of low tremulous or 'gurgling 1 notes, resembling somewhat 
the notes uttered by Rana boyhi" (T. I. Storer, Calif., 1925, p. 238). 

Breeding: This species breeds from January to March. The eggs are laid 



418 HANDBOOK OF FROGS AND TOADS 

in overflow areas of permanent pools, the mass attached to vegetation. The 
jelly is soft and viscid; the outline of individual eggs is evident on the sur- 
face. The egg is Yi2 inch (1.8-2.1 mm.), the envelopes %, %o, % inch (3.5 
mm., 4.4 mm., 8.5 mm.), the eggs black and white. The dark brownish, 
mottled tadpole is 3% inches (83 mm.) and has tooth ridges %. The larval 
period is 5-7 months. (Data from Storer, 1925.) The tadpoles transform from 
May to August at %-i% inches (18-30 mm.). 

This frog is one of earliest to spawn in California, the crest coming in Jan- 
uary and February. The customary size at transformation is probably 
22-30 mm. Of seven specimens taken by us Aug. 14, 1917, at Oceanside, Calif., 
none was smaller than 32 mm. and all were past transformation. In one set, 
that of the University of Michigan (no. 65203), appear nine transforming frogs 
25-35 mm. in body length, with tail stubs from 13-48 mm. Two specimens 
just transformed are 31 and 32 mm. and 14 specimens are 29-37 mm. 

In egg measurements one is not sure whether or not some other author 
interpreted the vitelline membrane as an envelope. We measured some of 
Storer's eggs from Thornhill, Calif., to be able to compare his statements with 
other material. We found eggs of i.8-2.i-mm. vitelh, a vitelline membrane, 
and three envelopes, the outer of which was loose and the two inner much 
firmer. We therefore agree with Storer's diagrams and measurements. 

Journal notes: Feb. 14, 1942, Hoover Ranch. Cook caught me an adult fe- 
male R. a. draytonii. 

March 10, Stanford. Anita Daugherty yesterday gave us R. a. draytonii eggs 
laid in lab from frogs taken at Los Trancos Creek March i. 

April 12. Out with Tracy and Ruth Storer. . . . After going along stream 
tributary to N. Fork Cosumnes River north of Plymouth, Calif., near the 
Amador-El Dorado County line, in the depressions of gold workings, saw a 
R. a. draytonii jump into a deep pool. It rested on the bottom. Brought net 
in front of it and poked it from rear as I pushed forward with net. Caught 
this one and three more. One was in grass; it leaped into a clump. Put net 
between clump and water. Then kicked from rear and in jumped my frog. 
These ugly deposits are from dredging or placer workings. 

April 22. With Leo T. Hadsall and party stopped beside drainage ditch on 
level, near Fresno, Calif.; some three or four R. a. draytonii jumped in. 

May 5, La Posta. At a creek beside a tule area saw some amphibian jump 
in. Had to know whether it was Bttfo calif ornictts, my search, or Rana a. 
draytomi. It proved the latter. Three and one-half to 4 miles south of Buck- 
man's Spring at bridge below confluence of La Posta and Cottonwood Cr. 
took one R. a. draytontt. Saw no Rana tadpoles. 

Authorities 9 corner: 
M. C. Dickerson, Gen., 1906, p. 216. 
T. I. Storer, Calif., 1925, p. 245. 
J. M. Linsdale, Nev., 1938, p. 25. 



FROGS: RANIDAE 419 

California Yellow-legged Frog, Thick-skinned Frog, Boyle's Frog 
Rana boylii boylii (Baird). Plate LXXXIX; Map 28. 

Range: Western Oregon, northern and central portions of California, 
chiefly west of the high Sierra Nevada. 

"Life zones Upper Sonoran and Transition" (J, Grinnell and C. L. Camp, 
Calif., 1917, p. 146). 

"Except for a small area in southwestern Oregon, the range of Rana boylii 
boylii jeems to lie entirely within the state of California. It includes the north- 
western part of the state, east to the McCloud River, Shasta County, and to 
the western foothills of the Sierra Nevada below 4100 feet altitude. The form 
occurs also at Mono Lake; in the Sierras it has been taken south to the vicinity 
of Walker Pass, Kern County; but where it meets the range of R. b. muscosa 
along the coast is not known. Specimens from the vicinity of Walker Pass 
show a more contrasted pattern of coloration, possibly indicating approach 
to the southern subspecies, described below" (C. L. Camp, Calif., 1917, p. 118). 

Habitat: Margins of springs, streams, and fresh water lakes (J. Grinnell and 
C. L. Camp, Calif., 1917); gravelly streams (K. Gordon, Ore., 1939). 

"It is confined to the immediate vicinity of permanent streams, at least those 
where water holes persist through the dry season. It is most common along 
streams having rocky beds, but occurs also in ones having mud buttoms" (H. 
S. Fitch, Ore., 1936, p. 640). 

"This little frog inhabits the slow flowing streams of the coastal areas, and 
may be found in considerable numbers in the semipermanent pools, formed 
as the streams become low at the end of the rainy season" (J. R. Slevin, B.C., 
1928, p. 139). 

At Stevens Creek near Palo Alto, Calif., in shallow and quiet areas we 
found small 1?. boylii. Up one or two little runs bordered by Rhits (poison 
oak), saw four more. 

T. I. Storer says: "The adult frogs of this species spend much of their time 
perched on rocks in the stream or on the bank, but in the latter place they 
never go more than two or three feet from the margin of the water. If ap- 
proached, even from the direction of the stream, they invariably seek safety 
by leaping into the water, and immediately swim with swift strokes down 
to the bottom. In streams with silt on the bottom they hide in the mud and 
silt which their movements stir up; in clear waters they take refuge under 
overhanging rocks." 

Size: Adults, i%~3 inches. Males, 39-67 mm. Females, 40-75 mm. It is the 
smallest of the three subspecies of Rana boylii. 

General appearance: This is a small frog with stout, broad body. The skin 
on the back, legs, and tympanum is thick and rough with small brownish 
papillae. The color of the back is black, light gray, greenish, or brownish with 
indistinct dark mottling. There is a patch of lighter color on top of the head 



4 20 



HANDBOOK OF FROGS AND TOADS 



with a darker area behind crossing the upper eyelids. The tympanic region 
is darker than the head. Red is never present in the coloration. The venter is 
white with pale yellow on the posterior part and on the hind legs. The throat 
and sides of the body are mottled with black. 

Of four adult specimens of R. b. 
boylii taken April 2, 1942, only one, a 
male, had a tendency to be spotted in 
the groin. All had spotted throats. 
Three males had short, light tubercular 
ridges back of the angle of the mouth. 
It was either a continuous fold or two 
to four tubercles in a row; then came a 
hiatus and then another ridge or patch 
of one to three tubercles. The skin was 
rougher than in Rana pretiosa or R. a. 
aurora. Quite often they had a red line 
along the dorsolateral fold, the lower 
belly lumiere green, underside of femur 
mustard yellow, then primuline yellow, 
then yellow ocher, becoming in rear 
portion aniline yellow. 

Color: Female. Mill Valley, Muir 
Woods, Calif., from John Needham, 
March 18, 1930. Dorsal color in general 
is buffy olive, light brownish olive, 
citrine-drab, or olive lake, coming to be, 
in the intervals between the leg and 
arm crossbars, Isabella color, clay color, 
or honey yellow on the tarsus and outer 
edge of the foot. The crossbars are Sac- 
cardo's olive. On rear of femur and in 
the groin are several rather prominent 
black, olive, or deep olive areas. On the 
rear of femur the dark areas have ecru- 
olive or light yellowish olive intervals. A ridge from rear of upper labial margin 
around angle of mouth, a short area back of eye and irregularly along back 
where dorsolateral fold would be, and patches on either side of knee have 
orange-cinnamon tubercles. The front of the tibia bears an empire yellow line 
separating dorsal from ventral surface. The top of the head from front of one 
eyelid to the other and forward on snout is Isabella color or old gold. Ventral 
surfaces: The central rear half of the femur is papillate, orange, or deep chrome, 
bounded behind by a more or less prominent black or deep olive band. In front 
of this orange area and on lower belly is a wash of picric yellow. Some of same 




Map 28 



FROGS: RANIDAE 



421 



color on groin, on rear of tibia, on venter of arm, and in axil of arm. The top of 
the foot has a wash of empire yellow. The throat and breast are white to pale 
pinkish buff with orange-cinnamon spots on throat and vaguely outlined on 
breast. These become verona brown spots on edge of lower jaw. Along side 
as back merges into belly are picric yellow spots among the deep olive. The 
eye is black, almost obscured by numerous specks of buff-pink or vinaceous- 
pink on lower portion giving place to light green-yellow and pale orange- 
yellow specks on the upper half. Pupil rim in lower half partakes of color 
of specks of that part, and so also the rim on the upper part. 

Structure: Head bioad and pointed; web of foot large, only slightly scal- 
loped; toes blunt, tips slightly expanded; thumb of male enlarged, the swelling 
having two lobes; tympanum small, distinct. 

"Rana boyht, Baird. A broad depressed ridge of skin on each side of back. 
Skin finely tubercular above. Head broader than long. Tympanum scarcely 
evident, pustulated. Tibia more than half the length of body; hind foot less 
than half this length; webbed entirely to the horny tips; outer toe decidedly 
longer than the third. An elongated tubercle at base of inner toe, with another 
opposite to it. Above dull reddish olivaceous, with indistinct blotches on the 
back, and fascia on the legs. Beneath yellowish, mottled anteriorly. Two inches 
long. Hab. California (interior)" (S. F. Baird, Gen., 1856, p. 62). 

"Those characters which pertain to this subspecies alone are: hind leg long, 
inside angle of bent tarsus reaching at least to narcs and usually beyond when 
leg is advanced along body; tibia elongate, reaching usually beyond anus when 
flexed and held at right angles to axis of body; fourth toe on reflexed hind foot 
never reaching beyond end of knee and often not quite to fold of skin below 
knee; head broad and pointed when viewed from above, its width two and 
one-third to two and two-thirds times in body-length; skin on back, legs and 
tympanum, thick and rough with minute brownish spines; color dorsally 
varying from nearly uniform to black to light gray, greenish or brownish, with 
darker markings, if present, usually indistinct; there is always a patch of 
lighter color on top of head between nares and eyes, and behind this a 
darker area crossing posterior half of each eyelid and merging insensibly be- 
hind into the general dorsal coloration" (C. L. Camp, Calif., 1917, pp. 117- 
118). 

"Size small; head broader than long, depressed; snout rounded, projecting 
beyond mouth; canthus rostralis distinct; loreal region concave; nostrils nearer 
to tip of snout than to orbit; distance between nostrils equals interorbital width. 
Interorbital width less than width of upper eyelid. Tympanum small, distinct, 
covered with small tubercles, about one-half diameter of eye. Fore limbs mod- 
erately robust; digits rather long, first and second equal, third much the long- 
est; no lateral fold along sides of fingers; subarticular tubercles small or 
moderately large, rounded, prominent; outer metatarsal tubercles elongate, 
prominent, inner metatarsal tubercle rounded, somewhat obscure; web full, 



422 HANDBOOK OF FROGS AND TOADS 

extending to tip of longest toe. Skin rugose, covered with warts or tubercles 
on back and sides; posterior surface of the thigh covered with small tubercles; 
dorso-lateral fold obscure; sacral hump rather prominent; a well developed 
fold from lip to side of neck or shoulder. Vomerine teeth in two oblique series, 
widely separated anteriorly, between and a little behind the choanae" (J. R. 
Slevin, B.C., 1928, p. 137). 

Voice: No description of the call is recorded. The frog has internal vocal 
sacs. 

Breeding: This frog breeds from the latter part of March to the first of May. 
The egg mass, in shallow water toward the margin of streams, is attached to 
sides of stones in the stream bed and is like a compact cluster of grapes, the 
individual eggs on the surface distinct, the jelly firm. The eggs are more 
closely set than are those in a R. ptpiens mass. 

"Three envelopes present. Outer envelope 3.88-4.47 mm., ave. 4.0 mm,; jelly 
firm. Middle envelope 2.58-3.35 mm., ave. 2.8 mm. Inner envelope 2.32-2.94 
mm., ave. 2.5 mm. All envelopes distinct. Vitellus 1.93-2.48 mm., average 2.2 
mm.; black above and white below. Mass 2 by 2 by i% to 2 by 4 by 2% inches 
in dimensions. Eggs firmly attached to each other. 909-1037 eggs in a mass. 
Deposited in shallow water near the margins of streams, attached to stones 
in stream bed. Pick up much sediment" (R. L. Livezey and A. H. Wright, 
Gen., 1947). 

The tadpole is medium, 2 inches (50 mm.), deep olive in color, and with 
tooth ridges %,%,% After 3 to 4 months, the tadpoles transform from June 
to September, at %-iYs inches (20-30 mm.). 

"Evans Creek (Oregon) on April 29, 1934. Numerous egg masses at- 
tached to large pebbles and newly hatched tadpoles were seen. The spawning 
places were in pools four or five inches deep where slow currents or eddies 
caused constant circulation of water around the egg masses" (H. S. Fitch, 
Ore., 1936, p. 640). 

A frog taken by J. C. Bradley, June 1907, Blue Lake, Humboldt Co., Calif., 
was 29 mm. and past transformation; and in a scries of six from Marin Co. 
by J. C. Bradley, which were 22.5-30 mm., none had arm scars. Transforma- 
tion of some of this group must have come at 20 mm. or smaller. H. C. Kellers, 
April 13, 1913, secured two specimens (USNM nos. 49874-5) 25 and 31 mm. 
These doubtless wintered over from the previous year. This same collector 
has taken this species in Calistoga, Napa Co., Calif., as small as 23 mm. Mr. 
Rutter, Oct. 17, 1899, in Mariposa, Calif., secured two specimens (nos. 
38830-31) 29.0 and 25.5 mm. These might be of the season. The specimens 
(USNM nos. 20895-20903) secured by H. W. Henshaw Oct. 17, 1893, at Ukiah, 
Mendocino Co., Calif., ranging from 20-29 mm -> ma Y a ls be of the same 
season. 

April 8, 1942. Ate at Trail, Ore. Went up the creek. In a spring area saw 
several R. boylii boylii. Took in side pools small R. b. boylii. They swim out 



FROGS: RANIDAE 423 

into deep water and then generally back to edge or to shallow water. In the 
middle of stream riffles saw an adult R. b. boylii swimming. While waiting 
for Bert to search above the bridge for more snakes (Thamnophis o. hydro- 
phila) I went to stream side near bridge. Across the stream I saw what I 
thought an adult R. b. boylii. This was a crossing place in the stream with flat 
rocks; just above were rapids and just below, deep pools with rapid current 
in water falling into pools. I waded across stream catching sight of two adult 
frogs. Then in a shallow side pocket with water flowing through but not 
rapidly, attached to a broken notch, was a mass of R. b. boyln eggs. The water 
was 3 to 4 inches deep. The egg cluster was somewhat plinthlike. Eggs were 
fresh and looked black. Bert came down and reported that this must be type 
locality for Fitch's T. o. hydrophila. No end of adult Rana b. boylii leaped into 
midstream ahead of him. It is interesting to see them helplessly carried over 
the shallow rock bottom by the current until they find a deep groove. They 
are shy frogs and you can't get within catching distance before they dash for 
the bottom under the edge of a near-by stone if there is a handy one. 

Journal notes: Feb. 14, 1942. Trip to Theodore Hoover ranch, Calif. Went 
up steep side of small creek and in it Rodgers secured a yellow-legged frog. 
Around sawmill in little shallow pools found several small R. boylii boylii. 
During the day the boys got me one or two more. 

March 15. Trip north. In Tuttle Ranch, Calif., took R. b. boylii. 

March 18. Started at 10:30 along Smith River road toward Smith River 
village, Calif. It is the east segment of the triangle. Along a bank covered with 
ferns and oxalis in a rill where water goes under a road, Anna caught a male 
R. b. boylii. In pool at other end of pipe I took two more. 

March 23. With Kenneth Gordon and group to Santiam River, Ore. In 
ponds beside stream under rock, Storm and Livezcy took three R. boylii boylti. 
Under board, Anna found a female. 

April 12. Out with Tracy and Ruth Storer. Region of dredging and placer 
working, north of Plymouth, Calif. Over in the stream Tracy and Ruth Storer 
and Anna kept seeing yellow-legged frogs leap into the water. This is a creek 
with plenty of nlHy spots. In a fast-water stretch Tracy Storer found a mass 
of R. b. boylii eggs attached to stone 3 or 4 feet from bank in water 10 inches 
deep, a plinth. Before we got through we found five more masses one very 
fresh and hardly expanded, this less plinthlike. 

April 1 8. Went to Hasting's Natural History Reservation to sec Jean Lins- 
dale. He has H. regtlla and R. boylii there. 

Authorities 9 corner: 
C. E. and M. D. Burt, Ariz., 1929, p. H. S. Fitch, Ore., 1936, pp. 640-641. 

432. H. S. Fitch, Ore., 1938, p. 148. 

J. Grinnell, J. J. Dixson, J. M. Linsdale, 

Calif., 1930, p. 143. 

"There is thus a good chance that on the desert slope of the San Gabriel 



414 HANDBOOK OF FROGS AND TOADS 

Mountains the frogs are boylii rather than muscosa' (C. L, Camp, Calif., 1917, 
p. 120). [A statement verified by Marr 26 years later.] 

"In a herpetological collection made along the San Gabriel River, San 
Gabriel Mountains, Los Angeles County, August n, 1940, I have four speci- 
mens of Rana boylii boylii and none of Rana boylii muscosa. ... It is barely 
possible that this frog has been introduced by some chance into the San Gabriel 
River" (}. C. Marr, Calif., 1943, p. 56). 

Sierra Madre Yellow-legged Frog, Southern Yellow-legged Frog 

Rana boylii muscosa Camp. Plate XC; Map 28. 

Range: San Gabriel, San Bernardino, and San Jacinto Mountains of south- 
ern California. "This form inhabits stream canons in southern California 
south of Tehachapi Pass. Localities of record extend from Pacoima Canon 
near San Fernando eastward and southward along the southern and western 
slopes of the San Gabriel, San Bernardino and San Jacinto ranges as far as 
Keen's Camp, in the latter range. It has been found also at Little Rock Creek 
on the north slope of the San Gabriel Mountains. The range in altitude is from 
1200 feet near Sierra Madrc to 6500 feet on Fish Creek in the San Bernardino 
range" (C. L. Camp, Calif., 1917^ p. 119). 

"Southern California south o Ventura County" (J. R. Slevin, Calif., 1934, 

p. 4). 

"The previously recorded southern limit of this species has been Keen 
Camp in the San Jacinto Mountains, Riverside County, California. Recently 
it has been found to be plentiful in Pauma Creek, where it flows through 
Upper Doane Valley, Palomar Mountain, San Diego County, at elevation 5100 
feet. The first specimens at this location were taken by Dr. E. H. Taylor of 
the University of Kansas during a collecting trip in which the writer partici- 
pated" (L. M. Klauber, Calif., 1929, pp. 15-16). 

Habitat: "Occupies the upper Sonoran and Transition life /ones. Lives 
along streams in narrow rockwalled canons" (J. Grinnell and C. L. Camp, 
Calif., 1917, p. 148). "More or less common in streams throughout (Los 
Angeles) County" (C. M. Bogert, Calif., 1930, p. 5). 

Size: Adults, i %~3/4 inches. Males, 44-66 mm. Females, 47-85 mm. Our 
material taken in Santa Anita Canyon and Snow Creek in spring, 1942, falls 
into three classes: young 33-40 mm., males 49-56 mm., and females 49- 
64 mm. 

General appearance: "Like Rana boylii boylti, but attaining much larger 
size, and (except in young) with no light patch in front of dark areas across 
upper eyelids. Dorsal ground color usually lighter than in R. b. boylti, light 
yellow to brown, contrasting with the darker mosslike patches on the back. 
Tips of toes more expanded than in boylii 9 (C. L. Camp, Calif., 1917, p. 119). 

Color: Male. San Gabriel Canyon, Los Angeles region, Calif., from Berry 



FROGS: RANIDAE 



4*5 




Plate LXXXIX. Rana boylii boylii. i. Plate XC. Rana boylii muscosa 

Male (X%). 2,3. Females (X%). 1,2- Females. 3,4. Males. 



426 HANDBOOK OF FROGS AND TOADS 

Campbell, April n, 1930. Upper parts from tip of snout back between eyes 
and lateral folds to vent ecru-olive or deep olive-buff. Costal folds dark olive- 
buff. Interspaces on hind legs dark olive-buff instead of ecru-olive. Color below 
costal fold olive-buff or ivory yellow and same on under parts and front half 
of under part of hind legs. An oblique spot from eye to angle of mouth, one 
from tympanum obliquely backwards, colonial buff. On front of femur, top 
of foot, axilla, slightly on head and front edge of arm deep colonial buff. 
Throat cartridge buff with fine buffy olive spottings. Spot on top of head and 
back brownish olive. Same color also on hind legs and forelegs, on sides be- 
coming buffy olive. Underside of head and hind ventral half of femur pale 
grayish vinaceous or pale vinaceous-fawn. Thumb excrescences clove brown, 
also underside of hand. 

Female. Upper Doane Valley, Palomar Mt., San Diego Co., Calif., from 
L. M. Klauber and Dr. E. H. Taylor, Sept. 2, 1928. Upper parts from tip of 
snout back between eyes and two costal folds to vent grayish olive or citrine- 
drab (lighter than in male). Costal fold dark to deep olive-buff, color below 
costal fold pale olive-buff, and the same on under parts. An oblique spot from 
eye to angle of mouth, one back of angle, one over arm insertion and one in 
axilla primrose yellow or olive-buff. Under parts of legs like venter. Where 
lighter color joins upper color considerable of deep colonial buff or chamois. 
This color comes strongly into light intervals between crossbars on hind legs 
and somewhat on forelegs. Some wash of same color on throat, which is not 
as spotted as in male. Sides below costal folds heavily spotted with brownish 
olive. Spots on dorsum from tip of snout to vent smaller and often more or 
less olive-ocher encircled. Dark line from eye to nostril to labial margin. Black 
spot or bar below eye. Edge of lower jaw black or brownish olive and colonial 
buff or pale olive-buff. Top of fingers deep colonial buff or chamois. Dark 
bars of hind legs brownish olive or olive with some grayish olive spots in the 
dark color. Hind webs dark with some chamois. Horizontal bar of black or 
olive through eye; vertical bar of same below pupil; interspaces between the 
horizontal and vertical, isabella color or even primrose yellow; upper portion 
of eye colonial buff with black dot at the top. Sometimes at forward edge of 
eye occurs pale yellow-green. 

Structure: Tips of toes more expanded than in R. b. boylii; dorsolateral fold 
indistinct, not pitted anteriorly; tympanum and area surrounding it very 
rough, covered with small tubercles; web of hind foot extending nearly to tips 
of toes, broader in expanse than in R. b. boyhi; swelling on thumb of male has 
two lobes, the constriction being diagonal not transverse. 

"Vomerine teeth on two oblique ridges between nares; head pointed in out- 
line as viewed from above, broad, its width entering body-length two and 
two-thirds times; hind limbs long, posterior side of bent tarsus reaching for- 
ward to snout; fourth toe on hind foot reaching forward not quite to knee- 
fold; dorsolateral fold indistinct, not pitted anteriorly; tympanum and area 



FROGS: RANIDAE 427 

surrounding it very rough, beset with small tubercles, web of hind foot ex- 
tending nearly to tips of toes; outer and inner metatarsal tubercles distinct; 
plantar tubercles very large; tips of toes expanded, disc-shaped; distal end of 
flexed tibia held at right angles to body reaching anus" (C. L. Camp, Calif., 
1917, p. 119). 

"Ground color above light gray or yellowish, with numerous black blotches 
or reticulation on head, body and limbs. Posterior surface of thigh rich yellow. 
Under surfaces yellowish or whitish, gular region slightly spotted or marbled 
with gray" (J. R. Slevin, Gen., 1928, p. 141). 

Voice: Nothing on record. 

Breeding: This species breeds during April. The eggs are not yet described. 
The tadpole is much like that of R. boylii sierrae and reaches 61 mm. in length. 
Its body is rather flat, the tail musculature heavy and almost uniform in width 
for about half the length of tail. Tooth ridges are variable, for example, % in 
one, % in another, /4 in another. Possibly they winter as tadpoles and trans- 
form in the spring. They transform at about % inch (20-24 mm.). 

Two tadpoles, collected Sept. 5, 1922, in San Jacinto Mountains, Dark 
Valley, by Prof. J. C. Bradley are 41 and 43 mm., not fully grown. The crests 
remain about the same to the point where musculature tapers. 

One specimen from near Clarcmont, Calif. , 1907, is 30 mm., just beyond 
transformation. The two smallest in the National Museum (nos. 54899- 
54900), taken by E. J. Brown in San Gabriel Canyon, Los Angeles Co., Calif., 
in February, 1916, are 24 and 26 mm. Apparently they were transformed the 
season before. This seems to imply transformation in the neighborhood of 
20 mm. 

On June 20, 1928, Storer secured at BlufI Lake, 3400 feet elevation, San 
Bernardino Mts. and Co., a tadpole, 61 mm. long, body 23 mm. Its tail was 
rounded; apparently the tadpole was approaching transformation; its tooth 
formula was % Other specimens had tails that were not broad, crests quite 
spotted. Some were practically transformed at 23 and 24 mm. 

On May i, 1909, C. L. Camp found specimens 20-28 mm.; on Feb. 17, 1918, 
J. Shaw took a 25-mm. specimen and on May 30, 1918, a 29-0101.; on Nov. 13, 
1937, W. F. Wood took specimens from 24-36 mm. One wonders if they winter 
over as tadpoles. 

Journal notes: April 26, 1942, Calif. Started for Santa Anita Canyon about 
9:30 A.M., hazy over the mountains. Rode about 3 miles up canyon to public 
parking spot. Went down trail to the river, about % mile down, and then 
a mile or so up the river. Near where trail went to ranger cabin crossed a 
very rocky, steep gorge with small amount of swift water. It was dark and 
shady at this point. We saw no frogs there. As we reached the river another 
side stream came in, White Creek. We went slowly up the river. At one spot 
beside the river but not connected with it was a pool, a rather scummy, dark- 
looking pool, small, perhaps 4 by 6 feet. Here Bert caught one male and one 



428 HANDBOOK OF FROGS AND TOADS 

small Rana b. muscosa and some newts, Triturus torosus, and found quantities 
of beautiful newt egg clusters attached to sticks. More were under a board and 
some were attached to a board lying at the end of the pool. (Caught two 
H. aremcolor.} Much granitic rock here light gray in color almost no sedi- 
ment in the stream. On our return, Bert stopped at White Creek and there 
caught three female R. b. muscosa , nice large ones. He carried them in a large 
wet bag. He had two newts with them. When we reached the car he put the 
bag into the thermos bottle with a little water. Either too much water or too 
much newt killed two of the females by the next day. A male JR. b. muscosa 
leaped into a quiet algae-covered pool filled with brush. Took young 
R. b. muscosa in this pool. In a quiet cutofl saw a female leap into creek and 
go under a stone. I was too leisurely about it. Lost it. I was angry to lose the 
female; saw no more except two or three young. 

On return came to Winter Creek. Here on a sandy bar was one female. It 
leaped into a quiet pool near the base of a falls. Another halfway up on falls 
leaped in and lodged under a stone part way down. Caught it. In a pool at 
top of this sloping granitic falls took another female in brushy edge. Many 
newts mating in this creek. A few trout in Winter Creek. 

April 30. Trip to Idyllwyld. At mouth of Strawberry Creek in cattail area 
in one quiet pool saw two Rana b. muscosa. Took one in tules below bridge, 
and later above bridge there leaped in from a boulder a female H. aremcolor. 
Went to Idyllwyld no luck. Tried several places. 

May 9, Palm Springs. Went up Snow Creek 5 miles. Here road ends where 
water company's possessions are. The Sierra Club of Los Angeles was meeting 
here. Some eight to ten people were along the creek awaiting their leader's 
coming. A beautiful cool mountain stream. There used to be a fish hatchery 
in the stream. At once back of car 1850 feet elevation we saw R. boylit 
muscosa; the first near weeds at edge leaped among some rocks but came up, 
as they frequently do, and was easy of capture. 

May 9, Snow Creek. These frogs were most frequently seated on the rocks. 
I would flop the net flat on the rock and the frog would leap into the netting. 
Sometimes they disappeared before I reached close enough to catch them. 
Occasionally I could approach close and seize them with my left hand. We saw 
only two or three small ones. I collected about five females before I got a male. 
In the end there were seven females and two males in the lot. A fine grayish- 
white, bouldery, tumbling stream of cold water. No wonder these muscosa 
are much lighter than the sterrae and boyhi. They are a shade-loving form and 
have a decided odor. They sat on damp rocks above the water line. The first 
male was darker than the rest. 

Authorities 9 corner: 
C. L. Camp, Calif., 1917, pp. 119-120. 
L. M. Klauber, Calif., 19343, p. 7. 



FROGS: RANIDAE 429 

Sierra Nevada Yellow-legged Frog, Western Frog, Pacific Frog, Stink Frog 

Rana boylii sierrae Camp. Plate XCI; Map 28. 

Range: California. Southern half of Sierra Nevada, above 7000 feet altitude 
from Yosemite National Park on the north to southern Tulare County on 
the south. 

Habitat: Meadows, streams, and lakes from 7000 to 10,500 feet in Yosemite 
Park, and to 11,500 feet near Mount Whitney. 

"In some of the lakes it was found in great numbers, appearing as soon 
as the ice had melted in late June. The tadpoles were at this time of large size 
and must have been hatched from eggs ot a laying not more recent than the 
previous year" (C. L. Camp, Calif., 1917!}, p. 122). 

Size: Adults, i%~3% inches. Males, 44-72 mm. Females, 48-85 mm. The 
measurements of 19 males and 30 females in the California Academy, Stanford 
University, and University of California collections gave a range of 44-65 mm. 
for males and 52-85 mm. for females; average male 55.5, mean 56 mm.; aver- 
age female 60 mm., mean 59, 66, 70 mm. 

General appearance: This frog is like Rana boyln boyln, but lacks the light 
patch on the head. The back is slightly roughened, is dark olive in color with 
some darker spots showing when it is wet. The under parts arc white on the 
throat, creamy or buff on the belly and the underside of the legs. The throat 
is marked with black spots. The sides are light, heavily mottled with distinct 
black blotches. The jaw is heavily mottled with light and dark. There is a 
distinct dark blotch on the upper part of the forearm. The inner surface of 
the arms is light, outer surface greenish, marked with dark blotches. The legs 
are marked with dark spots. The angle of the jaw forms a swollen light area 
marked with one or two dark spots, and back of this is another light tubercle 
or swollen area. The head is short and broad with the nostrils far apart, snout 
short, eyes small and widely separated. Costal folds low and inconspicuous. 
The young look something like newly transformed green frogs. 

"With the general characters of Rana boyln boylii, but hind leg usually 
shorter and head relatively narrower; tympanum smoother; and light patch 
on top of head wanting" (C. L. Camp, Calif., igi7b, p. 120). 

Color: Female. Yosemite Park, from C. A. Harwell, Oct. 12, 1929. (The 
frog was cold when described, hence dark.) Back deep olive, citrine-drab, 
light brownish olive, few black or dark spots centered with old gold between 
lateral folds from arm insertion level to vent. More heavily spotted on sides 
below lateral folds and spots sometimes light-centered there where belly color 
and spots are reticulate. Something of same spotting on front of dorsum of 
femur and rear of forelimb. Throat and upper and lower labial edge heavily 
spotted with black. A very light white or cartridge buff line begins below eye 
and extends across angle of mouth or just above it to arm insertion. Some- 
times this is interrupted below tympanum, then resumed diagonally down 



430 HANDBOOK OF FROGS AND TOADS 

from tympanum, then interrupted to appear as a swollen patch above arm 
insertion. The posttympanal and arm insertion cartridge buff patches may 
have two or three fine black spots on them and considerable black extending 
upward between them from arm insertion. Upper jaw and face pale olive- 
buff with numerous black spots. Tympanum clear, like background of dorsum 
or top of head. Background on hind legs and forelegs dark olive-buff or old 
gold; mottled or spotted with black or olive; lateral fold of same color. Under- 
side of hind legs and forelegs pale chalcedony yellow, cream color, or seafoam 
yellow; a tinge of same color on lower belly. On tarsus old gold or aniline 
yellow. Throat and pectoral region white. Underside of forefeet and hind feet 
light brownish drab or hair brown; iris black dotted with citron to honey 
yellow, or strontium yellow; pupil rim broken in front and behind. 

Male. Lake Alpine, 7300 ft., Alpine Co., Calif., from Tracy I. Storer, July 
29, 1930. Back buffy olive, citrine-drab, or grayish olive. Under the lens it may 
look deep olive-buff. On sides interspace color is deep olive-buff, ivory yellow, 
or cartridge buff, becoming chamois on edges of belly. The interspaces of 
hind legs and of forelegs are light brownish olive or almost Isabella color. 
Dorsolateral fold in front half isabella color or cinnamon-brown to tawny 
or sayal brown. Large spots of bister or auburn. Whole of interspaces and 
spots covered by spaced black points. On sides, dorsolateral fold, and top of 
femur these surmount larger papillae. Forearm, front of femur, rear of tibia, 
tarsus, and foot with transverse bars of same color as large dorsal spots. Lower 
rear half of femur with many papillae tipped with reed yellow or deep colonial 
buff. Throat to line of arm insertion and upper and lower labial fold heavily 
spotted with deep olive or citrine-drab. Middle belly reed yellow, its sides 
chamois. Underside of hind legs and forelegs colonial buff or cream-buff. In- 
side of tarsus and lower end of tibia chamois. Rear half of venter of femur 
vinaceous-drab or deep brownish vinaceous, each papilla tipped with deep 
olive-buff or, better, reed yellow or deep colonial buff. Underside of fingers 
and of toes like rear half of venter of femur. Cream buff or colonial buff fold 
from below eye to angle of mouth broken below tympanum, broken again, 
resumed as spot above arm insertion. Iris rim picric yellow or lemon yellow 
above. Iris mars orange or English red. 

Structure: Hind legs shorter; head relatively narrower; tympanum 
smoother than in Rana b. boyhi; web of hind foot very large, extending to 
tips of toes; tips of toes not much expanded; dorsolateral fold strongly pitted 
anteriorly; swelling on thumb of male bilobate. 

"Vomerine teeth rudimentary, on two oblique ridges nearly meeting be- 
tween and slightly behind nares . . . ; head viewed from above rounded in 
outline; head-width contained three times in body-length; hind limbs short, 
posterior side of bent tarsus reaching forward to anterior corner of eye; fourth 
toe on hind foot reaching forward to end of bent knee; dorso-Iateral fold 
indistinct, strongly pitted anteriorly; tympanum nearly smooth, with scat- 



FROGS: RANIDAE 431 

tered hispid points; web of hind foot very large, extending to tips of toes; 
outer metatarsal tubercle rudimentary, inner one small; plantar tubercles 
small; tips of toes not much expanded; distal end of flexed tibia, held at right 
angles to body, just reaching anus; color above, dark yellowish brown, ob- 
scurely marked with indefinite darker vermiculations; lower surface yellow, 
faintly dotted with brown beneath chin; upper lip below eye mottled; no 
dark cheek patch; hind limbs not distinctly barred with dark bands. 

"This seems to be the subspecies of boyln which approaches most closely 
to the species pretiosa" (C. L. Camp, Calit., 1917^ pp. 120-122). 

Voice: No data. No vocal sacs. Croaks not very loud in captive males. 

Breeding: As soon as ice has melted in the mountain lakes, these frogs breed 
in June and July. The eggs are approximately like those of Rana b. boylii (T. I. 
Storer, Calif., 1925). The tadpole is fairly large, 2% inches (72 mm.), its body 
flattened, the tail musculature wide for an inch or more, then, suddenly ta- 
pering, the tail tip rounded 01 spatulate. The crests are broader toward the tip 
than at the body. The tooth ridges are %, %, or %. The period of develop- 
ment is i year, the animal passing the winter as a tadpole. It transforms, dur- 
ing July and August, %-i % 2 inches (20-27 mm.) . 

In the summer of 1907, Prof. J. C. Bradley at Giant Forest, Calif., took 
transforming young measuring 21-26 mm. On Aug. 22-27, 1917, we took at 
Alta Meadows, Sequoia National Park, a series of transforming frogs which 
measured from 21 mm. to 27 mm., the average and mode being 23 mm. Of 
our twelve females, three had ripe eggs to be laid. 

At Davis, Calif., we examined some of Professor Storer's material. He had 
males 54-72 mm. and females 58-82 mm. in length. He had a young specimen 
36 mm. long July 14, and others, near transformation, 20, 21.5, 22, 22 mm. 
July 3, 1930. One tadpole of 54 mm. (body length 21.5 mm.) had a tail that 
tended to be rounded, even spatulate, at the tip. Its teeth were %. Those of 
another were %. The eggs had three jelly envelopes; the vitellus was 2.0 mm.; 
and the egg was about 7 mm. across. Egg masses were attached to the banks of 
a small creek. 

"One mass of eggs was collected by Osgorxl R. Smith on April 23, 1942 at 
Convict Creek, Mono County, California, at an elevation of 7,000 feet. This 
clutch was attached to stems of sedge and contained about 120 eggs. . . . 

"Mass plinth-like, about 28 X 40 mm., individual eggs one-quarter to one- 
half inch apart. There are besides the vitellinc capsule three gelatinous en- 
velopes, all clear and transparent. The outline of the egg is distinct and the firm 
jelly does not collect much debris. All envelopes arc quite distinguishable in 
good light without a lens, as is the vitelline capsule. Least in thickness is the 
middle envelope, the next in thickness is the inner, being about two times the 
thickness of the middle, and the thickest is the outer which is approximately 
twice that of the inner. The vitellus is black at the animal pole and grey-tan 
at the vegetal. Measurements are: vitellus 2.17 mm. (range 1.81 to 2.30 mm.); 



432 HANDBOOK OF FROGS AND TOADS 

vitclline capsule 2.58 mm. (range 2.25 to 2.75 mm.) ; inner envelope 3.93 mm. 
(range 3.75 to 4.81 mm.) ; middle envelope 4.60 mm. (range 4.25 to 5.00 mm.) ; 
outer envelope 7.19 mm. (range 6.43 to 7.87 mm.). 

"Measurements on the eggs preserved before they had time to attain their 
full size showed the vitellus to average somewhat less (av. 1.88 mm.) than 
the other eggs taken from natural surroundings" (R. L. Livezey and A. H. 
Wright, Gen., 1945, p. 703). 

Journal notes: Aug. 23, 1917, Alta Meadows, Calif. In the pools on the mats 
were plenty of Rana boylii sierrae. They stink like minks or other weasels. 
They are